Interbreeding between archaic and modern humans

6768:"Non-African populations (Han Chinese in Beijing and Utah residents with northern and western European ancestry) also show analogous patterns in the CSFS, suggesting that a component of archaic ancestry was shared before the split of African and non-African populations...One interpretation of the recent time of introgression that we document is that archaic forms persisted in Africa until fairly recently. Alternately, the archaic population could have introgressed earlier into a modern human population, which then subsequently interbred with the ancestors of the populations that we have analyzed here. The models that we have explored here are not mutually exclusive, and it is plausible that the history of African populations includes genetic contributions from multiple divergent populations, as evidenced by the large effective population size associated with the introgressing archaic population...Given the uncertainty in our estimates of the time of introgression, we wondered whether jointly analyzing the CSFS from both the CEU (Utah residents with Northern and Western European ancestry) and YRI genomes could provide additional resolution. Under model C, we simulated introgression before and after the split between African and non-African populations and observed qualitative differences between the two models in the high-frequency–derived allele bins of the CSFS in African and non-African populations (fig. S40). Using ABC to jointly fit the high-frequency–derived allele bins of the CSFS in CEU and YRI (defined as greater than 50% frequency), we find that the lower limit on the 95% credible interval of the introgression time is older than the simulated split between CEU and YRI (2800 versus 2155 generations B.P.), indicating that at least part of the archaic lineages seen in the YRI are also shared with the CEU..." 1897:(Pleistocene New Guinea and Australia), at least 44,000 years ago. It has also been observed that the fraction of Near Oceanian ancestry in Southeast Asians is proportional to the Denisovan admixture, except in the Philippines where there is a higher proportional Denisovan admixture to Near Oceanian ancestry. Reich et al. (2011) suggested a possible model of an early eastward migration wave of modern humans, some who were Philippine/New Guinean/Australian common ancestors that interbred with Denisovans, respectively followed by divergence of the Philippine early ancestors, interbreeding between the New Guinean and Australian early ancestors with a part of the same early-migration population that did not experience Denisovan gene flow, and interbreeding between the Philippine early ancestors with a part of the population from a much-later eastward migration wave (the other part of the migrating population would become East Asians). 378:(Spain), and Vindija Neanderthals indicates that of these three lineages, only the El Sidrón and Vindija Neanderthals display significant rates of gene flow (0.3–2.6%) into modern humans, suggesting that the El Sidrón and Vindija Neanderthals are more closely related than the Altai Neanderthal to the Neanderthals that interbred with modern humans about 47,000–65,000 years ago. Conversely, significant rates of modern human gene flow into Neanderthals occurred—of the three examined lineages—for only the Altai Neanderthal (0.1–2.1%), suggesting that modern human gene flow into Neanderthals mainly took place after the separation of the Altai Neanderthals from the El Sidrón and Vindija Neanderthals that occurred roughly 110,000 years ago. The findings show that the source of modern human gene flow into Neanderthals originated from a population of 33: 2001:

of DNA from 4 fossils found at Shum Laka in Cameroon dating from 8,000 to 3,000 BP. These individuals were found to derive most of their DNA from Central African hunter gatherers (Pygmy ancestors) and did not share the archaic DNA found in the Yoruba and Mande. The pattern of differences between Eastern, Central and Southern hunter gatherers when compared to the West African groups which had been found by Hammer was confirmed. In a second study Lipson et al. (2020) studied DNA extracted from 6 additional Eastern and Southcentral African fossils from the last 18,000 years. It was determined that their genetic origins could be accounted for by DNA contributions from Southern, Central and Eastern hunter gatherers, and that none of them had the archaic DNA found in the Yoruba.

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thought to have no Neanderthal admixture and were therefore used as reference samples. Thus, any overlap in Neanderthal admixture with Africans resulted in an underestimation of Neanderthal admixture in non-Africans and especially in Europeans. The authors give a single pulse of Neanderthal admixture after the out-of-Africa dispersal as the most parsimonious explanation for the enrichment in East Asians, but they add that variation in Neanderthal ancestry may also be attributed to dilution to account for the now-more-modest differences found. As a proportion of the total amount of Neanderthal sequence for each population, 7.2% of the sequence in Europeans is shared exclusively with Africans, while 2% of the sequence in East Asians is shared exclusively with Africans.

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HPD: 26,000–28,000), admixture fraction of 0.09 (95% HPD: 0.04–0.17), admixture time of 3,000 generations ago (95% HPD: 2,800–3,400), and an effective population size of 19,700 individuals (95% HPD: 19,300–20,200). We find that the lower bound of the admixture time is further back than the simulated split between CEU and YRI (2155 generations ago), providing some evidence in favor of a pre-Out-of-Africa event. This model suggests that many populations outside of Africa should also contain haplotypes from this introgression event, though detection is difficult because many methods use unadmixed outgroups to detect introgressed haplotypes (5, 53, 22). It is also possible that some of these haplotypes were lost during the Out-of-Africa bottleneck."

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offspring of Neanderthal mothers were raised in Neanderthal groups and became extinct with them, or that female Neanderthals and male Sapiens did not produce fertile offspring. However, the hypothesized incompatibility between Neanderthals and modern humans is contested by findings that suggest that the Y chromosome of Neanderthals was replaced by an extinct lineage of the modern human Y chromosome, which introgressed into Neanderthals between 100,000 and 370,000 years ago. Furthermore, the study concludes that the replacement of the Y chromosomes and mitochondrial DNA in Neanderthals after gene flow from modern humans is highly plausible given the increased

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to the studies of Skoglund and Lipson with ancient African DNA, the study also finds that at least part of this proposed archaic admixture is also present in Eurasians/non-Africans, and that the admixture event or events range from 0 to 124 ka B.P, which includes the period before the Out-of-Africa migration and prior to the African/Eurasian split (thus affecting in part the common ancestors of both Africans and Eurasians/non-Africans). Another recent study, which discovered substantial amounts of previously undescribed human genetic variation, also found ancestral genetic variation in Africans that predates modern humans and was lost in most non-Africans.

1761:(Western Galilee, Israel) and dated to 54.7±5.5 kyr BP, represents the first fossil evidence from the period when modern humans successfully migrated out of Africa and colonized Eurasia. It also provides the first fossil evidence that modern humans inhabited the southern Levant during the Middle to Upper Palaeolithic interface, contemporaneously with the Neanderthals and close to the probable interbreeding event. The morphological features suggest that the Manot population may be closely related to or may have given rise to the first modern humans who later successfully colonized Europe to establish early Upper Palaeolithic populations. 2535:"An analysis of archaic sequences in modern populations identifies ancestral genetic variation in African populations that likely predates modern humans and has been lost in most non-African populations...We found small amounts of Neanderthal ancestry in West African genomes, most likely reflecting Eurasian admixture. Despite their very low levels or absence of archaic ancestry, African populations share many Neanderthal and Denisovan variants that are absent from Eurasia, reflecting how a larger proportion of the ancestral human variation has been maintained in Africa." 1905:(i.e. America, Central Asia, East Asia, and South Asia). The researchers also made the surprising finding that South Asian populations display an elevated Denisovan admixture (when compared to other non-Oceanian populations with Denisovan ancestry), albeit the highest estimate (which are found in Sherpas) is still ten times lower than in Papuans. They suggest two possible explanations: There was a single Denisovan introgression event that was followed by dilution to different extents or at least three distinct pulses of Denisovan introgressions must have occurred. 1993:

not included in the pre-agricultural Eastern African hunter-gatherers, Southern African hunter-gatherer populations, or the genetic gradation between them. The West African groups carrying the archaic DNA include Yoruba from coastal Nigeria and Mende from Sierra Leon indicating that the ancient DNA was acquired long before the spread of agriculture and likely well before the Holocene (starting 11,600 BP), Such an archaic lineage must have separated before the divergence of San ancestors, which is estimated to have begun on the order of 200–300 thousand years ago.

219: 1881:(e.g. Aboriginal Australians, Near Oceanians, Polynesians, Fijians, eastern Indonesians, Philippine Mamanwa and Manobo), but not in certain western and continental Southeast Asian populations (e.g. western Indonesians, Malaysian Jehai, Andaman Onge, and mainland Asians), indicating that the Denisovan admixture event happened in Southeast Asia itself rather than mainland Eurasia. The observation of high Denisovan admixture in Oceania and the lack thereof in mainland Asia suggests that early modern humans and Denisovans had interbred east of the 290:

ancestry of East Asians, thus favoring more-complex models involving additional pulses of admixture between Neanderthals and the ancestors of East Asians. Such models show a pulse to ancestral Eurasians, followed by separation and an additional pulse to ancestral East Asians. It is observed that there is a small but significant variation of Neanderthal admixture rates within European populations, but no significant variation within East Asian populations. Prüfer et al. (2017) remarked that East Asians carry more Neanderthal DNA (2.3–2.6%) than

332: 1670:. In conjunction with archaeological and fossil evidence, the gene flow is thought likely to have occurred somewhere in Western Eurasia, possibly the Middle East. Through another approach—using one genome each of a Neanderthal, Eurasian, African, and chimpanzee (outgroup), and dividing it into non-recombining short sequence blocks—to estimate genome-wide maximum-likelihood under different models, an ancient population sub-structure in Africa was ruled out and a Neanderthal admixture event was confirmed. 3752:

the highest European and Near Eastern component (~40%), have also the highest amount of Neandertal ancestry (~60–70%) (Figure 3). On the contrary, South Morocco that exhibits the highest Sub-Saharan component (~60%), shows the lowest Neandertal signal (20%). Interestingly, the analysis of the Tunisian and N-TUN populations shows a higher Neandertal ancestry component than any other North African population and at least the same (or even higher) as other Eurasian populations (100–138%) (Figure 3).

1877:, indicating that there was interaction between the early ancestors of Melanesians with Denisovans but that this interaction did not take place in the regions near southern Siberia, where as-of-yet the only Denisovan remains have been found. In addition, Aboriginal Australians also show a relative increased allele sharing with Denisovans, compared to Eurasians and African populations, consistent with the hypothesis of increased admixture between Denisovans and Melanesians. 318:(~60–70%); and lowest among North African populations with greater Sub-Saharan admixture, such as in South Morocco (20%). Quinto et al. (2012) therefore postulate that the presence of this Neanderthal genetic signal in Africa is not due to recent gene flow from Near Eastern or European populations since it is higher among populations bearing indigenous pre-Neolithic North African ancestry. Low but significant rates of Neanderthal admixture has also been observed for the 1785:. He strongly emphasised that all living humans are of mixed origins. He held that this would best fit observations, and challenged the widespread idea that Neanderthals were ape-like or inferior. Basing his argument primarily on cranial data, he noted that the Danes, like the Frisians and the Dutch, exhibit some Neanderthaloid characteristics, and felt it was reasonable to "assume something was inherited" and that Neanderthals "are among our ancestors". 1935:, China) of 40,000 years BP showed a Neanderthal contribution within the range of today's Eurasian modern humans, but it had no discernible Denisovan contribution. It is a distant relative to the ancestors of many Asian and Native American populations, but post-dated the divergence between Asians and Europeans. The lack of a Denisovan component in the Tianyuan individual suggests that the genetic contribution had been always scarce in the mainland. 1840: 7960: 1682:(Portugal) features traits that indicate Neanderthal interbreeding with modern humans dispersing into Iberia. Considering the dating of the burial remains (24,500 years BP) and the persistence of Neanderthal traits long after the transitional period from a Neanderthal to a modern human population in Iberia (28,000–30,000 years BP), the child may have been a descendant of an already heavily admixed population. 1721: 7984: 367:

that the Neanderthal component in non-African modern humans is more closely related to the Vindija and Mezmaiskaya Neanderthals than to the Altai Neanderthal. These results suggest that a majority of the admixture into modern humans came from Neanderthal populations that had diverged (about 80–100kya) from the Vindija and Mezmaiskaya Neanderthal lineages before the latter two diverged from each other.

1478:, Italy) was homozygous for an ancestral allele of microcephalin, thus providing no support that Neanderthals contributed the D allele to modern humans and also not excluding the possibility of a Neanderthal origin of the D allele. Green et al. (2010), having analyzed the Vindija Neanderthals, also could not confirm a Neanderthal origin of haplogroup D of the microcephalin gene. 302:

alleles with Neanderthals as do non-African populations, whereas sub-Saharan African groups are the only modern human populations that generally did not experience Neanderthal admixture. The Neanderthal genetic signal among North African populations was found to vary depending on the relative quantity of North African, European, Near Eastern and sub-Saharan ancestry. Using

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Americans. In contrast, Prüfer et al. (2013) found that mainland Asian and Native American populations may have a 0.2% Denisovan contribution, which is about twenty-five times lower than Oceanian populations. The manner of gene flow to these populations remains unknown. However, Wall et al. (2013) stated that they found no evidence for Denisovan admixture in East Asians.

7972: 1740:, not present in earlier humans except Neanderthals of the late Middle and Late Pleistocene, thus suggesting affinity with Neanderthals. Concluding from the Oase 1 mandible, there was apparently a significant craniofacial change of early modern humans from at least Europe, possibly due to some degree of admixture with Neanderthals. 2291:

Eurasian ancestry in Ethiopians ranges from 11%–12% in the Gumuz to 53%–57% in the Amhara. Neanderthal ancestry proportion in Africans is correlated with gene flow from Eurasians. For example, knowing that today Eurasians carry ~2% of Neanderthal ancestry, we observed that East Africans (Ethiopians)

2000:

In the archaic DNA differences found by Hammer, et al., the pygmies (of Central Africa) are grouped with the San (of Southern Africa) in contrast to the Yoruba (of West Africa). Further clarification of the presence of archaic DNA in current West African populations with the extraction and sequencing

1900:

Finding components of Denisovan introgression with differing relatedness to the sequenced Denisovan, Browning et al. (2018) suggested that at least two separate episodes of Denisovan admixture has occurred. Specifically, introgression from two distinct Denisovan populations is observed in East Asians

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variation of modern humans. Looking at heterozygous individuals (carrying both Neanderthal and modern human versions of a gene), the allele-specific expression of introgressed Neanderthal alleles was found to be significantly lower in the brain and testes relative to other tissues. In the brain, this

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until a sudden increase of growth rate around 5,000 to 3,500 years BP. They occur at very high frequencies among East Asian populations in contrast to other Eurasian populations (e.g. European and South Asian populations). The findings also suggests that this Neanderthal introgression occurred within

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meiosis, as well as brain size and functioning. There are signals of positive selection, as the result of adaptation to diverse habitats, in genes involved with variation in skin pigmentation and hair morphology. In the immune system, introgressed variants have heavily contributed to the diversity of

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generations. This low rate of interbreeding would account for the absence of Neanderthal mitochondrial DNA from the modern human gene pool as found in earlier studies, as the model estimates a probability of only 7% for a Neanderthal origin of both mitochondrial DNA and Y chromosome in modern humans.

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As shown in an interbreeding model produced by Neves and Serva (2012), the Neanderthal admixture in modern humans may have been caused by a very low rate of interbreeding between modern humans and Neanderthals, with the exchange of one pair of individuals between the two populations in about every 77

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A higher Neanderthal admixture was found in East Asians than in Europeans, which is estimated to be about 20% more introgression into East Asians. This could possibly be explained by the occurrence of further admixture events in the early ancestors of East Asians after the separation of Europeans and

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It has been found that 50% of the Neanderthal genome is present among people in India, and 41% has been found in Icelanders. Previously it was found that about 20% of the Neanderthal genome was found in modern Eurasians, but the figure was also estimated at a third. A 2023 study found an introgession

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from Neanderthals to modern humans after the migration out of Africa. They estimated the proportion of Neanderthal-derived ancestry to be 1–4% of the Eurasian genome. Prüfer et al. (2013) estimated the proportion to be 1.5–2.1% for non-Africans, Lohse and Frantz (2014) infer a higher rate of 3.4–7.3%

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We show that North African populations, like all non-African humans, also carry the signature of admixture with Neandertals, and that the real geographical limit for Neandertal admixture is between sub-Saharan groups and the rest our results show that Neandertal genomic traces do not mark a division

1992:

Ancient DNA Data from a ~4,500 BP Ethiopian highland individual, and from Southern (~2,300–1,300 BP), and Eastern and South-Central Africa (~8,100–400 BP) has clarified that some West Africa populations have small amounts of excess alleles best explained by an archaic source in West Africans that is

1960:

The Denisovan's two HLA-A (A*02 and A*11) and two HLA-C (C*15 and C*12:02) allotypes correspond to common alleles in modern humans, whereas one of the Denisovan's HLA-B allotype corresponds to a rare recombinant allele and the other is absent in modern humans. It is thought that these must have been

1923:

It has been shown that Eurasians have some but significantly lesser archaic-derived genetic material that overlaps with Denisovans, stemming from the fact that Denisovans are related to Neanderthals—who contributed to the Eurasian gene pool—rather than from interbreeding of Denisovans with the early

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African populations) also have Neanderthal admixture, with this Neanderthal admixture in African individuals accounting for 17 megabases, which is 0.3% of their genome. According to the authors, Africans gained their Neanderthal admixture predominantly from a back-migration by peoples (modern humans

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Neanderthals, a draft sequence of the Neanderthal genome was published and revealed that Neanderthals shared more alleles with Eurasian populations (e.g. French, Han Chinese, and Papua New Guinean) than with sub-Saharan African populations (e.g. Yoruba and San). According to the authors Green et al.

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The results of the f4 ancestry ratio test (Table 2 and Table S1) show that North African populations vary in the percentage of Neandertal inferred admixture, primarily depending on the amount of European or Near Eastern ancestry they present (Table 1). Populations like North Morocco and Egypt, with

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Neandertal's genetic signal is higher in populations with a local, pre-Neolithic North African ancestry. Therefore, the detected ancient admixture is not due to recent Near Eastern or European migrations. Sub-Saharan populations are the only ones not affected by the admixture event with Neandertals.

2025:

Previous studies identified more recent event of admixture. About 350,000 years ago a genome of an "erectus-like" creature was injected into the Denisovan lineage. With the separation time of about 2 Ma ago and interbreeding that happened 350 ka ago, the two populations involved were more distantly

2004:

According to a study published in 2020, there are indications that 2% to 19% (or about ≃6.6 and ≃7.0%) of the DNA of four West African populations may have come from an unknown archaic hominin which split from the ancestor of humans and Neanderthals between 360 kya to 1.02 mya. However, in contrast

1904:

Exploring derived alleles from Denisovans, Sankararaman et al. (2016) estimated that the date of Denisovan admixture was 44,000–54,000 years ago. They also determined that the Denisovan admixture was the greatest in Oceanian populations compared to other populations with observed Denisovan ancestry

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than recent gene flow, the observation may have been due to ancient population sub-structure in Africa, causing incomplete genetic homogenization within modern humans when Neanderthals diverged while early ancestors of Eurasians were still more closely related to Neanderthals than those of Africans

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of East Africa. After identifying African and non-African ancestry among the Maasai, it can be concluded that recent non-African modern human (post-Neanderthal) gene flow was the source of the contribution since around an estimated 30% of the Maasai genome can be traced to non-African introgression

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It was later determined by Chen et al. (2020) that East Asians have 8% more Neanderthal ancestry, revised from the previous reports of 20% more Neanderthal ancestry, compared to Europeans. This stems from the fact that Neanderthal ancestry shared with Africans had been masked, because Africans were

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Recovering signals of ghost archaic introgression in African populations", section "S8.2" "We simulated data using the same priors in Section S5.2, but computed the spectrum for both YRI and CEU . We found that the best fitting parameters were an archaic split time of 27,000 generations ago (95%

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levels to compensate for low oxygen levels—such as at high altitudes—but this also has the maladaption of increasing blood viscosity. The Denisovan-derived variant on the other hand limits this increase of hemoglobin levels, thus resulting in a better altitude adaption. The Denisovan-derived EPAS1

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Findings indicate that the Denisovan gene flow event happened to the common ancestors of Aboriginal Filipinos, Aboriginal Australians, and New Guineans. New Guineans and Australians have similar rates of Denisovan admixture, indicating that interbreeding took place prior to their common ancestors'

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when compared to other Eurasian-derived populations and Africans. It is estimated that 4% to 6% of the genome in Melanesians derives from Denisovans, while no Eurasians or Africans displayed contributions of the Denisovan genes. It has been observed that Denisovans contributed genes to Melanesians

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of derived D alleles) into modern humans from an archaic human population that separated 1.1 million years ago (based on the separation time between D and non-D alleles), consistent with the period when Neanderthals and modern humans co-existed and diverged respectively. The high frequency of

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In Eurasia, modern humans have adaptive sequences introgressed from archaic humans, which provided a source of advantageous genetic variants that are adapted to local environments and a reservoir for additional genetic variation. Adaptive introgression from Neanderthals has targeted genes involved

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Neanderthals (Croatia). By high-coverage sequencing the genome of a 50,000-year-old female Vindija Neanderthal fragment, it was later found that the Vindija and Mezmaiskaya Neanderthals did not seem to differ in the extent of their allele-sharing with modern humans. In this case, it was also found

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consistent with several independent admixture events in the subcontinent have been found. It is currently unknown who these archaic African hominins were. A 2020 paper found that "despite their very low levels or absence of archaic ancestry, African populations share many Neanderthal and Denisovan

2013:

Hominins' presence in Eurasia begun at least 2 million years BP. Genetic evidence shows that thousands of years later when lineages of Neandertals and Denisovans started to expand into Eurasia, the continent was still inhabited by descendants of these archaic hominins, and their genetic admixture

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suggested that many Europeans bore traces of Neanderthal ancestry, but associated Neanderthal characteristics with primitivism, writing that since they "belong to a stage in the development of the human species, antecedent to the differentiation of any of the existing races, we may expect to find

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for in modern humans. The distribution of the D allele of microcephalin is high outside Africa but low in sub-Saharan Africa, which further suggest that the admixture event happened in archaic Eurasian populations. This distribution difference between Africa and Eurasia suggests that the D allele

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in the ancestors of East Asians, due smaller effective population sizes as they migrated to East Asia. Studies simulating admixture models indicate that a reduced efficacy of purifying selection against Neanderthal alleles in East Asians could not account for the greater proportion of Neanderthal

1880:

Reich et al. (2011) produced evidence that the highest presence of Denisovan admixture is in Oceanian populations, followed by many Southeast Asian populations, and none in East Asian populations. There is significant Denisovan genetic material in eastern Southeast Asian and Oceanian populations

1956:

alleles, it has been suggested that HLA-B*73 introgressed from Denisovans into modern humans in western Asia due to the distribution pattern and divergence of HLA-B*73 from other HLA alleles. Even though HLA-B*73 is not present in the sequenced Denisovan genome, HLA-B*73 was shown to be closely

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has been found in modern humans. This suggests that successful Neanderthal admixture happened in pairings with Neanderthal males and modern human females. Possible hypotheses are that Neanderthal mitochondrial DNA had detrimental mutations that led to the extinction of carriers, that the hybrid

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Genomic analysis suggests that there is a global division in Neanderthal introgression between sub-Saharan African populations and other modern human groups (including North Africans) rather than between African and non-African populations. North African groups share a similar excess of derived

5523:. "Currently available genetic and archaeological evidence is supportive of a recent single origin of modern humans in East Africa. However, this is where the consensus on human settlement history ends, and considerable uncertainty clouds any more detailed aspect of human colonization history." 1447:

Researchers found Neanderthal introgression of 18 genes—several of which are related to UV-light adaptation—within the chromosome 3p21.31 region (HYAL region) of East Asians. The introgressive haplotypes were positively selected in only East Asian populations, rising steadily from 45,000 years

1481:

It has been found that HLA-A*02, A*26/*66, B*07, B*51, C*07:02, and C*16:02 of the immune system were contributed from Neanderthals to modern humans. After migrating out of Africa, modern humans encountered and interbred with archaic humans, which was advantageous for modern humans in rapidly

1888:

Skoglund and Jakobsson (2011) observed that particularly Oceanians, followed by Southeast Asians populations, have a high Denisovans admixture relative to other populations. Furthermore, they found possible low traces of Denisovan admixture in East Asians and no Denisovan admixture in Native

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Examining European modern humans in regards to the Altai Neanderthal genome in high-coverage, results show that Neanderthal admixture is associated with several changes in cranium and underlying brain morphology, suggesting changes in neurological function through Neanderthal-derived genetic

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We found that North African populations have a significant excess of derived alleles shared with Neandertals, when compared to sub-Saharan Africans. This excess is similar to that found in non-African humans, a fact that can be interpreted as a sign of Neandertal admixture. Furthermore, the

2021:

Roger et al. (2020) describes an event of admixture that occurred soon after Neandersovans (common ancestor of Neanderthals and Denisovans) started to expand into Eurasia. They met a lineage of superarchaic hominins that had been separated from African homo lineages since at least 2 Ma ago.

1419:

were found to have been introgressed from Neanderthals into modern humans (shown in East Asians and Europeans), suggesting that these genes gave a morphological adaptation in skin and hair to modern humans to cope with non-African environments. This is likewise for several genes involved in

1996:

The hypothesis that there has been archaic line in the ancestry of present-day Africans that originated before the San, Pygmies and East African hunter gatherers (and the Eurasians) is supported by a line of evidence independent from the Skoglund findings based on long haplotypes with deep

1505:

Furthermore, correlating the genotypes of introgressed Neanderthal alleles with the expression of nearby genes, it is found that archaic alleles contribute proportionally more to variation in expression than nonarchaic alleles. Neanderthal alleles affect expression of the immune genes

423:. They also contained relatively high numbers of genes specific to testes. This means that modern humans have relatively few Neanderthal genes that are located on the X chromosome or expressed in the testes, suggesting male infertility as a probable cause. It may be partly affected by 3746:

North African populations have a complex genetic background. In addition to an Indegenous genetic component, they exhibit signals of European, sub-Saharan and Near Eastern admixture as previously described Tunisian Berbers and Saharawi are those populations with highest North African

32: 1796:, according to which anatomically modern humans left Africa about 50,000 years ago and replaced Neanderthals with little or no interbreeding. Yet some scholars still argued for hybridisation with Neanderthals. The most vocal proponent of the hybridisation hypothesis was 186:

A 2016 paper in the journal Evolutionary Biology argued that introgression of DNA from other lineages enabled humanity to migrate to, and succeed in, numerous new environments, with the resulting hybridization being an essential force in the emergence of modern humans.

1943:

There are large genomic regions devoid of Denisovan-derived ancestry, partly explained by infertility of male hybrids, as suggested by the lower proportion of Denisovan-derived ancestry on X chromosomes and in genes that are expressed in the testes of modern humans.

1747:, falling anatomically largely in line with the earliest (Middle Paleolithic) African modern humans, also show traits that are distinctively Neanderthal, suggesting that a solely Middle Paleolithic modern human ancestry was unlikely for European early modern humans. 1788:

Carleton Stevens Coon in 1962 found it likely, based upon evidence from cranial data and material culture, that Neanderthal and Upper Paleolithic peoples either interbred or that the newcomers reworked Neanderthal implements "into their own kind of tools".

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carrying Neanderthal admixture) that had diverged from ancestral Europeans (postdating the split between East Asians and Europeans). This back-migration is proposed to have happened about 20,000 years ago. However, some scientists, such as geneticist

1689:(Romania) of 35,000 years BP shows a morphological pattern of European early modern humans, but possesses archaic or Neanderthal features, suggesting European early modern humans interbreeding with Neanderthals. These features include a large 6348:

Llorente, M.G.; Jones, E.R.; Eriksson, A.; Siska, V.; Arthur, K.W.; Arthur, J.W.; Curtis, M.C.; Stock, J.T.; Coltorti, M.; Pieruccini, P.; et al. (2015). "Ancient Ethiopian genome reveals extensive Eurasian admixture in Eastern Africa".

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spectrum, it was shown that the recent admixture model had the best fit to the results while the ancient population sub-structure model had no fit—demonstrating that the best model was a recent admixture event that was preceded by a

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Daniel Harris; Alexander Platt; Matthew E.B. Hansen; Shaohua Fan; Michael A. McQuillan; Thomas Nyambo; Sununguko Wata Mpoloka; Gaonyadiwe George Mokone; Gurja Belay; Charles Fokunang; Alfred K. Njamnshi; Sarah A. Tishkoff (2023).

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patterns, a recent admixture event is likewise confirmed by the data. From the extent of linkage disequilibrium, it was estimated that the last Neanderthal gene flow into early ancestors of Europeans occurred 47,000–65,000 years

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Hershkovitz, Israel; Marder, Ofer; Ayalon, Avner; Bar-Matthews, Miryam; Yasur, Gal; Boaretto, Elisabetta; et al. (28 January 2015). "Levantine cranium from Manot Cave (Israel) foreshadows the first European modern humans".

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event among modern humans – thus confirming recent admixture as the most parsimonious and plausible explanation for the observed excess of genetic similarities between modern non-African humans and Neanderthals. On the basis of

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Most of us alive today carry inside our cells at least some DNA from a species that last saw the light of day tens of thousands of years ago. And we all carry different bits—to the extent that if you could add them all up,

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in the left hemisphere, Neanderthal admixture is positively correlated with gray matter volume. The results also show evidence for a negative correlation between Neanderthal admixture and white matter volume in the

3814:

Furthermore, the Neandertal's genetic signal is higher in populations with a local, pre-Neolithic North African ancestry. Therefore, the detected ancient admixture is not due to recent Near Eastern or European

2468: 5780: 310:, where it was at the same level or even higher than that of Eurasian populations (100–138%); high among North African populations carrying greater European or Near Eastern admixture, such as groups in North 4822:

Ham, Ellen; et al. (2019). "The relative roles of maternal survival and inter-personal violence as selection pressures on the persistence of Neanderthal hypercoagulability alleles in modern Europeans".

2017:

Genetic studies show two major events of genetic admixture from superarchaics, suggesting that in the late middle Pleistocene, Eurasia was inhabited by at least two separate populations of ancient hominins.

434:

as a result of strong selection against deleterious Neanderthal alleles. The overlap of many deserts of Neanderthal and Denisovan sequences suggests that repeated loss of archaic DNA occur at specific loci.

1619:

A Neanderthal allele inherited by modern humans, SNP rs3917862, is with associated with hypercoagulability. This can be harmful, but women lacking the allele are 0.1% more likely to die in childbirth.

1769:

The interbreeding has been discussed ever since the discovery of Neanderthal remains in the 19th century, though earlier writers believed that Neanderthals were a direct ancestor of modern humans.

1957:

associated to the Denisovan-derived HLA-C*15:05 from the linkage disequilibrium. From phylogenetic analysis, however, it has been concluded that it is highly likely that HLA-B*73 was ancestral.

3126: 89:

Neanderthal-derived DNA has been found in the genomes of most or possibly all contemporary populations, varying noticeably by region. It accounts for 1–4% of modern genomes for people outside

1616:, Neanderthal genetic variants are found in highest frequency in genes expressed in the brain, whereas Denisovan DNA has the highest frequency in genes expressed in bones and other tissues. 1989:

Rapid decay of fossils in Sub-Saharan African environments makes it currently unfeasible to compare modern human admixture with reference samples of archaic Sub-Saharan African hominins.

7013: 441:

Consistent with the hypothesis that purifying selection has reduced Neanderthal contribution in present-day modern human genomes, Upper Paleolithic Eurasian modern humans (such as the

159:

and some Southeast Asian populations. An estimated 4–6% of the genome of modern Melanesians is derived from Denisovans, but the highest amounts detected thus far are found in the

2600:"Morning Person? You Might Have Neanderthal Genes to Thank. - Hundreds of genetic variants carried by Neanderthals and Denisovans are shared by people who like to get up early" 6780: 1533:

Studying the high-coverage female Vindija Neanderthal genome, Prüfer et al. (2017) identified several Neanderthal-derived gene variants, including those that affect levels of

346:

Presenting a high-quality genome sequence of a female Altai Neanderthal, it has been found that the Neanderthal component in non-African modern humans is more related to the

306:

statistical analysis, the Neanderthal inferred admixture was observed to be: highest among the North African populations with highest North African ancestry such as Tunisian

2163: 2460: 1961:

contributed from Denisovans to modern humans, because it is unlikely to have been preserved independently in both for so long due to HLA alleles' high mutation rate.

1901:(e.g. Japanese and Han Chinese), whereas South Asians (e.g. Telugu and Punjabi) and Oceanians (e.g. Papuans) display introgression from one Denisovan population. 36:

Map of western Eurasia showing areas and estimated dates of possible Neanderthal–modern human hybridization (in red) based on fossil samples from indicated sites.

6041: 3884: 1916:

in particular were found to possess the highest level of Denisovan ancestry in the world, with ~30%–40% more than even that found in Australians and Papuans (

86:

events into modern humans are estimated to have happened about 47,000–65,000 years ago with Neanderthals and about 44,000–54,000 years ago with Denisovans.

3621:

between African and non-Africans but rather a division between Sub-Saharan Africans and the rest of modern human groups, including those from North Africa.

5693: 1972:

gene variant, associated with hemoglobin concentration and response to hypoxia, for life at high altitudes from the Denisovans. The ancestral variant of

1237: 415:

Neanderthal contribution in modern humans due to negative selection, partly caused by hybrid male infertility. These regions were most-pronounced on the

7093: 2609: 1502:. This downregulation suggests that modern humans and Neanderthals possibly experienced a relative higher rate of divergence in these specific tissues. 1107: 3968:

Krings, M.; Stone, A.; Schmitz, R.W.; Krainitzki, H.; Stoneking, M.; Pääbo, Svante (1997). "Neandertal DNA Sequences and the Origin of Modern Humans".

7053: 3029: 7103: 3118: 1736:(Romania) of 34,000–36,000 C years BP presents a mosaic of modern, archaic, and possible Neanderthal features. It displays a lingual bridging of the 7945: 7789: 2961: 1127: 496: 5414: 5661: 7098: 1087: 8015: 7847: 6924:

Wandell, P. (2013). "Happy New Year Homo erectus? More evidence for interbreeding with archaics predating the modern human/Neanderthal split".

1793: 7108: 5111: 7852: 7730: 7005: 4787:

Akkuratov, Evgeny E; Gelfand, Mikhail S; Khrameeva, Ekaterina E (2018). "Neanderthal and Denisovan ancestry in Papuans: A functional study".

3523:"Selection and Reduced Population Size Cannot Explain Higher Amounts of Neandertal Ancestry in East Asian than in European Human Populations" 2068: 1569:. In regards to modern human brain morphology, Neanderthal admixture is positively correlated with an increase in sulcal depth for the right 259:, have doubts about how extensive the flow of DNA back to Africa would have been, finding the signal of Neanderthal admixture "really weak". 226: 1920:), suggesting that distinct Islander Denisovan populations existed in the Philippines which admixed with modern humans after their arrival. 171:, have none. In addition, low traces of Denisovan-derived ancestry have been found in mainland Asia, with an elevated Denisovan ancestry in 183:

variants that are absent from Eurasia, reflecting how a larger proportion of the ancestral human variation has been maintained in Africa."

1557:

variation. Neanderthal admixture is associated with an expansion of the posterolateral area of the modern human skull, extending from the

281:

East Asians, dilution of Neanderthal ancestry in Europeans by populations with low Neanderthal ancestry from later migrations, or reduced

1474:

originated from Neanderthals according to Lari et al. (2010), but they found that a Neanderthal individual from the Mezzena Rockshelter (

8025: 7809: 7918: 5487: 1801: 430:

Deserts of Neanderthal sequences may also be caused by genetic drift involving intense bottlenecks in the modern human population and

141: 1908:

A study in 2021 analyzing archaic ancestry in 118 Philippine ethnic groups discovered an independent admixture event into Philippine

438:

It has also been shown that Neanderthal ancestry has been selected against in conserved biological pathways, such as RNA processing.

7784: 4479:"Evidence that the adaptive allele of the brain size gene microcephalin introgressed into Homo sapiens from an archaic Homo lineage" 4175:

Petr, Martin; Hajdinjak, Mateja; Fu, Qiaomei; Essel, Elena; Rougier, Hélène; Crevecoeur, Isabelle; et al. (25 September 2020).

1706: 4176: 2153: 7928: 7824: 7705: 1613: 1460:, a critical regulatory gene for brain volume, originated from an archaic human population. The results show that haplogroup D 1277: 382:

from about 100,000 years ago, predating the out-of-Africa migration of the modern human ancestors of present-day non-Africans.

5630: 7923: 7224: 5322: 1981:

gene variant is common in Tibetans and was positively selected in their ancestors after they colonized the Tibetan plateau.

4978:"The early Upper Paleolithic human skeleton from the Abrigo do Lagar Velho (Portugal) and modern-human emergence in Iberia" 1997:

divergences from other human haplotypes including Lachance et al.(2012), Hammer et al., 2011, and Plagnol and Wall (2006).

194:

inherited by modern humans from Neanderthals and Denisovans may biologically influence the daily routine of modern humans.

7046: 1728:

skull (cast depicted), found in Peştera cu Oase, displays archaic traits due to possible hybridization with Neanderthals.

7794: 7671: 7557: 5594: 1874: 1421: 717: 489: 3215: 3144:

says you could reconstitute something like one-third of the Neanderthal genome and 90 per cent of the Denisovan genome.

7988: 7892: 7710: 7231: 7085: 6033: 1067: 466: 4111: 5601: 7976: 7750: 7681: 7217: 7210: 7203: 6451:

Scally, A.; Durban, R. (2012). "Revising the human mutation rate: implications for understanding human evolution".

6301:"Evolutionary History and Adaptation from High-Coverage Whole-Genome Sequences of Diverse African Hunter-Gatherers" 3880: 3764:

Sánchez-Quinto, F.; Botigué, L.R.; Civit, S.; Arenas, C.; Ávila-Arcos, M.C.; Bustamante, C.D.; et al. (2012).

3696:

Sánchez-Quinto, F.; Botigué, L.R.; Civit, S.; Arenas, C.; Ávila-Arcos, M.C.; Bustamante, C.D.; et al. (2012).

3633:

Sánchez-Quinto, F.; Botigué, L.R.; Civit, S.; Arenas, C.; Ávila-Arcos, M.C.; Bustamante, C.D.; et al. (2012).

3570:

Sánchez-Quinto, F.; Botigué, L.R.; Civit, S.; Arenas, C.; Ávila-Arcos, M.C.; Bustamante, C.D.; et al. (2012).

2599: 1777:

Until the early 1950s, most scholars thought Neanderthals were not in the ancestry of living humans. Nevertheless,

1170: 904: 896: 888: 331: 8020: 7933: 7829: 7725: 1425: 1008: 94: 6947:"Approximate Bayesian computation with deep learning supports a third archaic introgression in Asia and Oceania" 218: 7964: 7267: 7039: 5683: 2400: 1441: 1412:

immune genes, of which there's an enrichment of introgressed alleles that suggest a strong positive selection.

256: 4730:

Gregory, M.D.; Kippenhan, J.S.; Eisenberg, D.P.; Kohn, P.D.; Dickinson, D.; Mattay, V.S.; et al. (2017).

3911:

Kuhlwilm, M.; Gronau, I.; Hubisz, M.J.; de Filippo, C.; Prado-Martinez, J.; Kircher, M.; et al. (2016).

2154:"A handful of recent discoveries have shattered anthropologists' picture of where humans came from, and when" 7902: 7839: 7799: 7245: 3867:

Furthermore we find that the Maasai of East Africa have a small but significant fraction of Neanderthal DNA.

2039: 1534: 978: 482: 1456:

Evans et al. (2006) had previously suggested that a group of alleles collectively known as haplogroup D of

1407:

with keratin filaments, sugar metabolism, muscle contraction, body fat distribution, enamel thickness, and

8010: 7897: 7872: 7867: 7760: 7715: 7676: 7238: 4021:

Serre, D.; Langaney, A.; Chech, M.; Teschler-Nicola, M.; Paunovic, M.; Mennecier, P.; et al. (2004).

2073: 1743:

The earliest (before about 33 ka BP) European modern humans and the subsequent (Middle Upper Paleolithic)

1662: 1486: 937: 457: 17: 4976:

Duarte, C.; Maurício, J.; Pettitt, P.B.; Souto, P.; Trinkaus, E.; Plicht, H. van der; Zilhão, J. (1999).

2951: 7281: 6951: 5802:

Rasmussen, M.; Guo, X.; Wang, Y.; Lohmueller, K.E.; Rasmussen, S.; Albrechtsen, A.; et al. (2011).

5723: 5402: 1809: 1657: 1598: 709: 247:

in Eurasia. In 2017, Prüfer et al. revised their estimate to 1.8–2.6% for non-Africans outside Oceania.

4606:

Abi-Rached, L.; Jobin, M. J.; Kulkarni, S.; McWhinnie, A.; Dalva, K.; Gragert, L.; et al. (2011).

2826:

Prüfer, K.; de Filippo, C.; Grote, S.; Mafessoni, F.; Korlević, P.; Hajdinjak, M.; et al. (2017).

3158: 7862: 7765: 7666: 7627: 7572: 7477: 7181: 7174: 6960: 6877: 6813: 6731: 6674: 6617: 6507: 6252: 6188: 6128: 5986: 5928: 5815: 5738: 5544: 5372: 5271: 5210: 5151: 5053: 4989: 4927: 4743: 4619: 4549: 4490: 4431:"Neanderthal Introgression at Chromosome 3p21.31 was Under Positive Natural Selection in East Asians" 4318: 4259: 4191: 4146: 3924: 3777: 3709: 3646: 3583: 3428: 3362: 3230: 3073: 2839: 2719: 2653: 2558: 2416: 2199: 1917: 1817: 1686: 1606: 1594: 1570: 1542: 1346: 1147: 994: 863: 848: 431: 251: 208: 5870:

Reich, D.; Patterson, N.; Kircher, M.; Delfin, F.; Nandineni, M.R.; Pugach, I.; et al. (2011).

3349:

Sankararaman, S.; Mallick, S.; Dannemann, M.; Prüfer, K.; Kelso, J.; Pääbo, S.; et al. (2014).

7274: 6299:

Lachance, J.; Vernot, B.; Elbers, C.C.; Ferwerda, B.; Froment, A.; Bodo, J.M.; et al. (2012).

5626: 5107: 2984: 2706:

Prüfer, K.; Racimo, F.; Patterson, N.; Jay, F.; Sankararaman, S.; Sawyer, S.; et al. (2014) .

2243:"Chad Genetic Diversity Reveals an African History Marked by Multiple Holocene Eurasian Migrations" 1778: 1694: 1690: 1550: 1482:

restoring HLA diversity and acquiring new HLA variants that are better adapted to local pathogens.

1470: 379: 286: 7983: 6239:

Huerta-Sánchez, E.; Jin, X.; Asan; Bianba, Z.; Peter, B.M.; Vinckenbosch, N.; et al. (2014).

5722:

Reich, D.; Green, R.E.; Kircher, M.; Krause, J.; Patterson, N.; Durand, E.Y.; et al. (2010).

4307:"40,000-Year-Old Individual from Asia Provides Insight into Early Population Structure in Eurasia" 2640:

Green, R.E.; Krause, J.; Briggs, A.W.; Maricic, T.; Stenzel, U.; Kircher, M.; et al. (2010).

2461:"Ancient human relative interbred with ancestors of modern humans as recently as 50,000 years ago" 268:

from modern humans to Neanderthals around 250,000 years ago, and estimated that roughly 6% of the

7819: 7686: 7661: 7537: 6925: 6476: 6384: 5952: 5568: 5295: 5071: 4917: 4225: 4003: 3159:"Diverse African genomes reveal selection on ancient modern human introgressions in Neanderthals" 3099: 2956: 2929: 2604: 2574: 2223: 2014:

made its way into genome of Neanderthals and Denisovans and later indirectly into modern humans.

1737: 1257: 90: 79: 44: 4852:"Ancient Structure in Africa Unlikely to Explain Neanderthal and Non-African Genetic Similarity" 1733: 1581:

of the left hemisphere. Neanderthal admixture is also positively correlated with an increase in

1429: 445:) carry more Neanderthal DNA (about 4–5%) than present-day Eurasian modern humans (about 1–2%). 222: 167:. While some Southeast Asian Negrito populations carry Denisovan admixture, others, such as the 5653: 229:

laureate and one of the researchers who published the first sequence of the Neanderthal genome.

7938: 7804: 7745: 7735: 7506: 7470: 7324: 7118: 6986: 6903: 6839: 6759: 6700: 6643: 6586: 6535: 6468: 6433: 6376: 6330: 6278: 6216: 6154: 6097: 6014: 5944: 5901: 5841: 5772: 5560: 5514: 5462: 5287: 5238: 5179: 5089: 5017: 4955: 4881: 4804: 4769: 4701: 4645: 4577: 4518: 4452: 4404: 4344: 4305:

Yang, M.A.; Gao, X.; Theunert, C.; Tong, H.; Aximu-Petri, A.; Nickel, B.; et al. (2017).

4287: 4217: 4089: 4054: 3995: 3950: 3858: 3827:

Wall, J.D.; Yang, M.A.; Jay, F.; Kim, S.K.; Durand, E.Y.; Stevison, L.S.; et al. (2013).

3805: 3737: 3674: 3611: 3552: 3503: 3454: 3415:

Nielsen, R.; Akey, J.M.; Jakobsson, M.; Pritchard, J.K.; Tishkoff, S.; Willerslev, E. (2017).

3388: 3312: 3281:

Wall, J.D.; Yang, M.A.; Jay, F.; Kim, S.K.; Durand, E.Y.; Stevison, L.S.; et al. (2013).

3256: 3178: 3091: 3010: 2921: 2865: 2797: 2745: 2679: 2526: 2499: 2442: 2370: 2282: 2215: 2134: 2116: 1774:

them in the lowest of these races, all over the world, and in the early stages of all races".

1710: 1465: 391: 303: 291: 234: 137: 98: 48: 3214:

Meyer, M.; Kircher, M.; Gansauge, M.T.; Li, H.; Racimo, F.; Mallick, S.; et al. (2012).

136:

compared to both East Asians and Europeans. However, other research finds higher Neanderthal

7814: 7567: 7384: 7346: 7193: 6976: 6968: 6893: 6885: 6829: 6821: 6749: 6739: 6690: 6682: 6633: 6625: 6576: 6566: 6525: 6515: 6460: 6423: 6415: 6366: 6358: 6320: 6312: 6268: 6260: 6206: 6196: 6144: 6136: 6087: 6079: 6004: 5994: 5936: 5891: 5883: 5831: 5823: 5762: 5754: 5746: 5552: 5535: 5504: 5496: 5454: 5380: 5279: 5228: 5218: 5169: 5159: 5138:

Trinkaus E.; Moldovan O.; Milota S.; Bîlgăr A.; Sarcina L.; Athreya S.; et al. (2003).

5079: 5061: 5007: 4997: 4945: 4935: 4871: 4863: 4832: 4796: 4759: 4751: 4691: 4683: 4635: 4627: 4567: 4557: 4508: 4498: 4442: 4394: 4384: 4334: 4326: 4277: 4267: 4207: 4199: 4154: 4081: 4044: 4034: 3985: 3977: 3940: 3932: 3848: 3840: 3795: 3785: 3727: 3717: 3664: 3654: 3601: 3591: 3542: 3534: 3493: 3485: 3444: 3436: 3378: 3370: 3302: 3294: 3246: 3238: 3170: 3081: 3000: 2992: 2985:"50,000 years of Evolutionary History of India: Insights from ~2,700 Whole Genome Sequences" 2911: 2855: 2847: 2787: 2779: 2735: 2727: 2669: 2661: 2566: 2516: 2508: 2432: 2424: 2360: 2350: 2272: 2262: 2254: 2207: 2158: 2124: 2106: 1754: 1702: 1652: 929: 878: 448:

Rates of selection against Neanderthal sequences varied for European and Asian populations.

347: 125: 4608:"The Shaping of Modern Human Immune Systems by Multiregional Admixture with Archaic Humans" 2768:"Neandertal Admixture in Eurasia Confirmed by Maximum-Likelihood Analysis of Three Genomes" 1839: 7621: 7594: 7492: 7450: 7062: 6784: 5872:"Denisova Admixture and the First Modern Human Dispersals into Southeast Asia and Oceania" 5605: 4373:"Evolutionary and Medical Consequences of Archaic Introgression into Modern Human Genomes" 3990: 3141: 1590: 1538: 962: 731: 687: 93:, although estimates vary, and either none or up to 0.3% for those in Sub-Saharan Africa. 4672:"Impacts of Neanderthal-Introgressed Sequences on the Landscape of Human Gene Expression" 6964: 6881: 6817: 6735: 6678: 6621: 6511: 6256: 6192: 6132: 5990: 5932: 5819: 5742: 5548: 5376: 5275: 5214: 5155: 5057: 4993: 4931: 4747: 4623: 4553: 4494: 4322: 4263: 4195: 4150: 3928: 3781: 3713: 3650: 3587: 3432: 3366: 3234: 3077: 3005: 2843: 2723: 2657: 2562: 2420: 2203: 1804:. Trinkaus claimed various fossils as products of hybridised populations, including the 7441: 7145: 7131: 7070: 6981: 6946: 6898: 6865: 6834: 6801: 6754: 6719: 6695: 6662: 6638: 6605: 6581: 6554: 6530: 6495: 6428: 6403: 6325: 6300: 6273: 6240: 6211: 6176: 6149: 6116: 6092: 6067: 6009: 5974: 5896: 5871: 5836: 5803: 5767: 5618: 5509: 5482: 5384: 5233: 5198: 5084: 5041: 4950: 4905: 4876: 4851: 4764: 4731: 4696: 4671: 4640: 4607: 4572: 4537: 4513: 4478: 4399: 4372: 4339: 4306: 4282: 4247: 3945: 3912: 3853: 3828: 3800: 3765: 3732: 3697: 3669: 3634: 3606: 3571: 3547: 3522: 3498: 3473: 3449: 3416: 3383: 3350: 3307: 3282: 3251: 2860: 2827: 2792: 2767: 2740: 2707: 2674: 2641: 2521: 2495:"Insights into human genetic variation and population history from 929 diverse genomes" 2494: 2437: 2404: 2365: 2338: 2277: 2242: 2129: 2094: 1714: 1667: 1558: 1449: 1437: 783: 770: 700: 674: 375: 351: 102: 5174: 5139: 4536:

Lari, M.; Rizzi, E.; Milani, L.; Corti, G.; Balsamo, C.; Vai, S.; et al. (2010).

4137:

Wang, C.C.; Farina, S.E.; Li, H. (2013) . "Neanderthal DNA and modern human origins".

4049: 4022: 3981: 1868:(e.g. Papua New Guinean and Bougainville Islander) share relatively more alleles with 8004: 7609: 7462: 7152: 6778: 6175:

Fu, Q.; Meyer, M.; Gao, X.; Stenzel, U.; Burbano, H.A.; Kelso, J.; Paabo, S. (2013).

5956: 5688: 5590: 5012: 4977: 4229: 4212: 2933: 2546: 2227: 2190:

Price, Michael (31 January 2020). "Africans, too, carry Neanderthal genetic legacy".

2063: 1834: 1805: 1797: 1770: 1698: 1578: 1566: 1562: 1546: 1499: 1461: 1457: 970: 371: 355: 319: 269: 110: 83: 6480: 6388: 5572: 4007: 3103: 2578: 2405:"The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans" 7615: 7603: 7545: 7419: 7367: 7332: 7257: 7164: 7006:"Artificial Intelligence Has Found an Unknown 'Ghost' Ancestor in The Human Genome" 6068:"Analysis of Human Sequence Data Reveals Two Pulses of Archaic Denisovan Admixture" 5299: 2900:"Identifying and Interpreting Apparent Neanderthal Ancestry in African Individuals" 2898:

Chen, Lu; Wolf, Aaron B.; Fu, Wenqing; Li, Liming; Akey, Joshua M. (January 2020).

2051: 1928: 1882: 1852: 1848: 1720: 1679: 1647: 1627: 1601: 1582: 1574: 1378: 952: 919: 833: 757: 744: 442: 424: 416: 396: 363: 339: 238: 172: 152: 75: 5441: 4248:"Extremely Rare Interbreeding Events Can Explain Neanderthal DNA in Living Humans" 6571: 5458: 5314: 4940: 4904:

Sankararaman, S.; Patterson, N.; Li, H.; Pääbo, S.; Reich, D; Akey, J.M. (2012).

4562: 4477:

Evans, P.D.; Mekel-Bobrov, N.; Vallender, E.J.; Hudson, R.R.; Lahn, B.T. (2006).

4272: 4158: 4039: 3790: 3722: 3659: 3596: 2355: 2211: 1530:, which can be specific to cell type and is influenced by environmental stimuli. 155:

and Europe. The highest rates, by far, of Denisovan admixture have been found in

7499: 7426: 7398: 7391: 7339: 3844: 3298: 2947: 2783: 2595: 2043: 1932: 1913: 1894: 1865: 1631: 1586: 1024: 648: 412: 335: 164: 133: 129: 117: 52: 6972: 6686: 6419: 6316: 6241:"Altitude adaptation in Tibetans caused by introgression of Denisovan-like DNA" 6083: 5919:

Cooper, A.; Stringer, C.B. (2013). "Did the Denisovans Cross Wallace's Line?".

5887: 5483:"A geographically explicit genetic model of worldwide human-settlement history" 4755: 4687: 3538: 3489: 2916: 2899: 2258: 7484: 7405: 6629: 6140: 6117:"Philippine Ayta possess the highest level of Denisovan ancestry in the world" 5654:"Life on the edge: was a Gibraltar cave last outpost of the lost neanderthal?" 4800: 4330: 4085: 3174: 2996: 2570: 2428: 1977: 1758: 1744: 1495: 1433: 250:

According to a later study by Chen et al. (2020), Africans (specifically, the

242:(2010), the observed excess of genetic similarity is best explained by recent 168: 106: 6494:

Hammer, M.F.; Woerner, A.E.; Mendez, F.L.; Watkins, J.C.; Wall, J.D. (2011).

5804:"An Aboriginal Australian Genome Reveals Separate Human Dispersals into Asia" 4732:"Neanderthal-Derived Genetic Variation Shapes Modern Human Cranium and Brain" 3766:"North African Populations Carry the Signature of Admixture with Neandertals" 3698:"North African Populations Carry the Signature of Admixture with Neandertals" 3635:"North African Populations Carry the Signature of Admixture with Neandertals" 3572:"North African Populations Carry the Signature of Admixture with Neandertals" 3182: 2120: 7551: 7456: 6606:"Ancient West African foragers in the context of African population history" 6520: 6362: 6201: 5999: 5940: 5827: 5223: 5164: 5066: 5002: 4867: 4631: 4503: 4447: 4430: 4203: 3242: 3086: 3061: 2851: 2665: 2512: 2111: 2047: 2042:(AI), that suggests the existence of an unknown human ancestor species, not 1869: 1844: 1635: 1490: 1362: 1193: 1039: 661: 243: 121: 56: 6990: 6907: 6889: 6843: 6825: 6763: 6744: 6704: 6663:"Ancient DNA and deep population structure in sub-Saharan African foragers" 6647: 6590: 6539: 6472: 6437: 6380: 6334: 6282: 6220: 6158: 6101: 6018: 5948: 5905: 5845: 5776: 5724:"Genetic history of an archaic hominin group from Denisova Cave in Siberia" 5564: 5518: 5466: 5344:

Boule, Marcellin (1911–1913). "L'homme fossile de La Chapelle-aux-Saints".

5291: 5242: 5183: 5093: 5021: 4959: 4885: 4808: 4773: 4705: 4649: 4581: 4522: 4456: 4408: 4348: 4291: 4221: 4093: 4058: 3954: 3862: 3809: 3741: 3678: 3615: 3556: 3507: 3458: 3392: 3316: 3260: 3095: 3014: 2925: 2869: 2801: 2749: 2683: 2530: 2446: 2374: 2286: 2219: 2138: 2026:

related than any pair of human populations previously known to interbreed.

6802:"Recovering signals of ghost archaic introgression in African populations" 6720:"Recovering signals of ghost archaic introgression in African populations" 4123: 4072:

Wall, J.D.; Hammer, M.F. (2006). "Archaic admixture in the human genome".

3999: 7412: 7377: 7354: 7075: 5598: 4389: 2267: 2035: 1953: 1909: 1813: 1475: 1324: 1302: 1213: 635: 420: 282: 156: 60: 51:. The interbreeding happened in several independent events that included 6371: 6264: 6066:

Browning, S.R.; Browning, B.L.; Zhou, Y.; Tucci, S.; Akey, J.M. (2018).

5750: 5283: 5075: 3936: 3829:"Higher Levels of Neanderthal Ancestry in East Asians than in Europeans" 3440: 3374: 3283:"Higher Levels of Neanderthal Ancestry in East Asians than in Europeans" 2731: 2708:"The complete genome sequence of a Neanderthal from the Altai Mountains" 147:

Denisovan-derived ancestry is largely absent from modern populations in

7562: 7138: 5758: 4836: 3030:"Genomes of modern Indian people include wide range of Neanderthal DNA" 1750: 1416: 1281: 818: 810: 796: 620: 359: 311: 307: 160: 3913:"Ancient gene flow from early modern humans into Eastern Neanderthals" 3216:"A High-Coverage Genome Sequence from an Archaic Denisovan Individual" 3062:"Resurrecting Surviving Neandertal Lineages from Modern Human Genomes" 6115:

Larena, Maximilian; McKenna, James; Sanchez-Quinto, Federico (2021).

5042:"Early modern humans from the Pestera Muierii, Baia de Fier, Romania" 4177:"The evolutionary history of Neanderthal and Denisovan Y chromosomes" 4023:"No Evidence of Neandertal mtDNA Contribution to Early Modern Humans" 3351:"The genomic landscape of Neanderthal ancestry in present-day humans" 1885:

that divides Southeast Asia according to Cooper and Stringer (2013).

1725: 1489:

effects in modern humans, contributing to the genomic complexity and

1453:

the ancestral population shared by East Asians and Native Americans.

1408: 179: 148: 67: 6464: 5556: 3474:"Complex History of Admixture between Modern Humans and Neandertals" 2952:"Neanderthal Genes Hint at Much Earlier Human Migration From Africa" 2545:

Rogers Ackermann, Rebecca; Mackay, Alex; Arnold, Michael L. (2016).

5500: 5110:. Smithsonian National Museum of Natural History. 23 January 2010. 7637: 7528: 7518: 7031: 6945:

Mondal, Mayukh; Bertranpedt, Jaume; Leo, Oscar (16 January 2019).

6930: 4922: 1973: 1969: 1965: 1838: 1719: 1527: 330: 315: 217: 31: 6177:"DNA analysis of an early modern human from Tianyuan Cave, China" 5533:

Stringer, Chris (June 2003). "Human evolution: Out of Ethiopia".

4906:"The Date of Interbreeding between Neandertals and Modern Humans" 2339:"Outstanding questions in the study of archaic hominin admixture" 7296: 6866:"Denisovan ancestors interbred with a distantly related hominin" 4850:

Yang, M.A.; Malaspinas, A.S.; Durand, E.Y.; Slatkin, M. (2012).

4538:"The Microcephalin Ancestral Allele in a Neanderthal Individual" 3119:"Who are you? How the story of human origins is being rewritten" 2828:"A high-coverage Neandertal genome from Vindija Cave in Croatia" 2095:"Midfacial Morphology and Neandertal–Modern Human Interbreeding" 2093:

Churchill, Steven E.; Keys, Kamryn; Ross, Ann H. (August 2022).

1623: 1523: 1519: 1515: 1511: 1507: 191: 71: 7035: 2493:

Bergström, A; McCarthy, S; Hui, R; Almarri, M; Ayub, Q (2020).

1638:

may biologically influence the daily routine of modern humans.

5199:"European early modern humans and the fate of the Neandertals" 1395: 461: 144:, than in Europeans (though not higher than in East Asians). 1685:

The remains of an early Upper Paleolithic modern human from

1792:

By the early 2000s, the majority of scholars supported the

6404:"Reconstructing Prehistoric African Population Structure" 5140:"An early modern human from the Peştera cu Oase, Romania" 78:, interbreeding between Neanderthals and Denisovans with 5040:

Soficaru, Andrei; Dobos, Adrian; Trinkaus, Erik (2006).

2399:

Sankararaman, Sriram; Mallick, Swapan; Patterson, Nick;

2292:

had ~1% Neanderthal ancestry and ~50% Eurasian ancestry.

1485:

It is found that introgressed Neanderthal genes exhibit

5481:

Liu H, Prugnolle F, Manica A, Balloux F (August 2006).

5363:

G.E. Smith (1928). "Neanderthal Man Not Our Ancestor".

419:, with fivefold lower Neanderthal ancestry compared to 1757:

of a modern human that was recently discovered at the

1469:

the D haplogroup (70%) suggest that it was positively

27:

Evidence of human hybridization during the Paleolithic

4429:

Ding, Q.; Hu, Y.; Xu, S.; Wang, J.; Jin, L. (2014) .

1420:

medical-relevant phenotypes, such as those affecting

109:), who derive a large portion of their ancestry from 6864:

Rogers, A.R.; Harris, N.S.; Achenbach, A.A. (2020).

3417:"Tracing the peopling of the world through genomics" 175:

populations compared to other mainland populations.

7911: 7885: 7838: 7776: 7695: 7654: 7647: 7593: 7526: 7517: 7440: 7365: 7312: 7295: 7255: 7191: 7162: 7117: 7084: 7069: 6555:"Possible ancestral structure in human populations" 5319:

Collected Essays: Volume VII, Man's Place in Nature

5035: 5033: 5031: 4789:

Journal of Bioinformatics and Computational Biology

1847:genome was sequenced from a fragment of the distal 7541:(archaic homo sapiens, anatomically modern humans) 6496:"Genetic evidence for archaic admixture in Africa" 5440: 2983:Elise Kerdoncuff; et al. (20 February 2024). 2893: 2891: 2889: 2887: 2885: 2883: 2881: 2879: 2034:In 2019, scientists discovered evidence, based on 128:. According to some research, it is also lower in 3209: 3207: 3205: 3203: 3201: 3199: 2488: 2486: 1678:The early Upper Paleolithic burial remains of a 6500:Proceedings of the National Academy of Sciences 6181:Proceedings of the National Academy of Sciences 5979:Proceedings of the National Academy of Sciences 5865: 5863: 5861: 5859: 5857: 5855: 5203:Proceedings of the National Academy of Sciences 5144:Proceedings of the National Academy of Sciences 5133: 5131: 5129: 5046:Proceedings of the National Academy of Sciences 4982:Proceedings of the National Academy of Sciences 4899: 4897: 4895: 4725: 4723: 4721: 4719: 4717: 4715: 4670:McCoy, R.C.; Wakefield, J.; Akey, J.M. (2017). 4483:Proceedings of the National Academy of Sciences 3276: 3274: 3272: 3270: 2590: 2588: 1713:to notch crest position, and a narrow scapular 41:Interbreeding between archaic and modern humans 6800:Durvasula, Arun; Sankararaman, Sriram (2020). 6294: 6292: 5595:Modern Humans, Neanderthals May Have Interbred 3906: 3904: 3902: 2701: 2699: 2697: 2695: 2693: 1855:(replica depicted) found in the Denisova cave. 1781:proposed interbreeding in 1907 in the article 7047: 6034:"Meet Your Ancient Relatives: The Denisovans" 4074:Current Opinion in Genetics & Development 3344: 3342: 3340: 3338: 3336: 3334: 3332: 3330: 3328: 3326: 2394: 2392: 2390: 2388: 2386: 2384: 1680:modern human child from Abrigo do Lagar Velho 490: 8: 6718:Arun Durvasula; Sriram Sankararaman (2020). 4472: 4470: 4468: 4466: 3410: 3408: 3406: 3404: 3402: 3055: 3053: 3051: 3049: 3047: 3045: 3043: 2821: 2819: 2817: 2815: 2813: 2811: 1732:The early modern human Oase 1 mandible from 6553:Plagnol, Vincent; Wall, Jeffrey D. (2006). 6234: 6232: 6230: 6061: 6059: 5968: 5966: 5717: 5715: 5713: 5711: 4665: 4663: 4661: 4659: 4366: 4364: 4362: 4360: 4358: 2761: 2759: 2642:"A Draft Sequence of the Neandertal Genome" 1622:In December 2023, scientists reported that 413:large genomic regions with strongly reduced 399:in Neanderthals relative to modern humans. 190:In December 2023, scientists reported that 7651: 7523: 7316: 7309: 7123: 7081: 7054: 7040: 7032: 6919: 6917: 6859: 6857: 6855: 6853: 6170: 6168: 5256: 5254: 5252: 4971: 4969: 4601: 4599: 4597: 4595: 4593: 4591: 4105: 4103: 2635: 2633: 2631: 2629: 2627: 497: 483: 18:Archaic human admixture with modern humans 6980: 6929: 6897: 6833: 6753: 6743: 6694: 6637: 6580: 6570: 6529: 6519: 6427: 6370: 6324: 6272: 6210: 6200: 6148: 6091: 6008: 5998: 5895: 5835: 5766: 5508: 5315:"The Aryan Question and Pre-Historic Man" 5232: 5222: 5173: 5163: 5083: 5065: 5011: 5001: 4949: 4939: 4921: 4875: 4763: 4695: 4639: 4571: 4561: 4512: 4502: 4446: 4424: 4422: 4420: 4418: 4398: 4388: 4338: 4281: 4271: 4211: 4048: 4038: 3989: 3944: 3852: 3799: 3789: 3731: 3721: 3668: 3658: 3605: 3595: 3546: 3497: 3448: 3382: 3306: 3250: 3085: 3004: 2915: 2859: 2791: 2739: 2673: 2520: 2436: 2364: 2354: 2332: 2330: 2328: 2326: 2324: 2322: 2320: 2276: 2266: 2185: 2183: 2181: 2128: 2110: 272:genome was inherited from modern humans. 7946:Human evolutionary developmental biology 6032:Flatow, I.; Reich, D. (31 August 2012). 5684:"Not a lasting last for the Neandertals" 4246:Neves, Armando; Serva, Maurizio (2012). 4241: 4239: 2318: 2316: 2314: 2312: 2310: 2308: 2306: 2304: 2302: 2300: 1964:Tibetan people received an advantageous 1709:, the relative perpendicular mandibular 4170: 4168: 2085: 6402:Skoglund, Pontus; et al. (2012). 5876:The American Journal of Human Genetics 3527:The American Journal of Human Genetics 3478:The American Journal of Human Genetics 2547:"The Hybrid Origin of "Modern" Humans" 1651:were to Neanderthals. On the basis of 116:Neanderthal-derived DNA is highest in 7731:Evolutionary models of human drug use 5975:"Archaic human ancestry in East Asia" 5696:from the original on 16 February 2020 5413:(in Danish). Royal Library, Denmark. 4371:Dolgova, O.; Lao, O. (18 July 2018). 2612:from the original on 14 December 2023 2471:from the original on 21 November 2022 2069:Multiregional origin of modern humans 7: 7971: 7004:Dockrill, Peter (11 February 2019). 5973:Skoglund, P.; Jakobsson, M. (2011). 5664:from the original on 20 January 2023 5593:193.2007, H. 2593 (3 March), 28–32. 3521:Kim, B.Y.; Lohmueller, K.E. (2015). 2964:from the original on 31 January 2020 1334: 1312: 1290: 1266: 1246: 1226: 1202: 1182: 1159: 1136: 1116: 1096: 1076: 1056: 113:, have ~1% Neanderthal-derived DNA. 3129:from the original on 25 August 2017 1929:modern human from the Tianyuan cave 7016:from the original on 23 April 2022 6661:Lipson, Mark; et al. (2022). 6604:Lipson, Mark; et al. (2020). 5488:American Journal of Human Genetics 5420:from the original on 16 March 2012 5405:[Race Studies in Denmark] 5385:10.1038/scientificamerican0828-112 2766:Lohse, K.; Frantz, L.A.F. (2014). 2166:from the original on 25 April 2022 2152:Woodward, Aylin (5 January 2020). 1802:Washington University in St. Louis 1537:and vitamin D, and that influence 386:Mitochondrial DNA and Y chromosome 338:Neanderthal skull reconstitution, 59:, as well as several unidentified 25: 5652:Sample, Ian (13 September 2006). 5633:from the original on 19 June 2010 5599:Humans and Neanderthals interbred 5325:from the original on 26 July 2011 4110:Mason, P.H.; Short, R.V. (2011). 3887:from the original on 11 June 2022 2337:Wolf, A. B.; Akey, J. M. (2018). 2241:Haber, Marc; et al. (2016). 1927:The skeletal remains of an early 7982: 7970: 7959: 7958: 6044:from the original on 2 July 2018 5783:from the original on 17 May 2020 3879:Bekker, Henk (23 October 2017). 427:of X chromosome genes in males. 323:from about 100 generations ago. 5439:Coon, Carleton Stevens (1962). 5114:from the original on 4 May 2018 4856:Molecular Biology and Evolution 4435:Molecular Biology and Evolution 3472:Vernot, B.; Akey, J.M. (2015). 3060:Vernot, B.; Akey, J.M. (2014). 3028:James Woodford (6 March 2024). 1464:37,000 years ago (based on the 5453:(3563). New York: Knopf: 208. 3991:11858/00-001M-0000-0025-0960-8 1952:Exploring the immune system's 1924:ancestors of those Eurasians. 1701:, an asymmetrical and shallow 1642:Population substructure theory 1597:. In the area overlapping the 1589:volume localized to the right 140:in Melanesians, as well as in 101:speaking populations from the 82:took place several times. The 1: 8016:Ancient human genetic history 3982:10.1016/S0092-8674(00)80310-4 3881:"Neues Museum in Berlin 1175" 1835:Australasians § Genetics 1541:, visceral fat accumulation, 233:On 7 May 2010, following the 6787:Supplementary Materials for 6572:10.1371/journal.pgen.0020105 5619:"The Lagar Velho 1 Skeleton" 5459:10.1126/science.140.3563.208 4941:10.1371/journal.pgen.1002947 4563:10.1371/journal.pone.0010648 4273:10.1371/journal.pone.0047076 4159:10.1016/j.quaint.2012.02.027 4040:10.1371/journal.pbio.0020057 3791:10.1371/journal.pone.0047765 3723:10.1371/journal.pone.0047765 3660:10.1371/journal.pone.0047765 3597:10.1371/journal.pone.0047765 2356:10.1371/journal.pgen.1007349 2212:10.1126/science.367.6477.497 1422:systemic lupus erythematosus 276:Subpopulation admixture rate 7989:Evolutionary biology Portal 5617:Foley, Jim (31 July 2000). 4112:"Neanderthal-human Hybrids" 3845:10.1534/genetics.112.148213 3299:10.1534/genetics.112.148213 2784:10.1534/genetics.114.162396 2009:Archaic hominins in Eurasia 1693:breadth, a relatively flat 1494:was most pronounced at the 390:No evidence of Neanderthal 8042: 8026:Anatomically modern humans 6973:10.1038/s41467-018-08089-7 6687:10.1038/s41586-022-04430-9 6420:10.1016/j.cell.2017.08.049 6317:10.1016/j.cell.2012.07.009 6084:10.1016/j.cell.2018.02.031 5888:10.1016/j.ajhg.2011.09.005 4756:10.1038/s41598-017-06587-0 4688:10.1016/j.cell.2017.01.038 3539:10.1016/j.ajhg.2014.12.029 3490:10.1016/j.ajhg.2015.01.006 2917:10.1016/j.cell.2020.01.012 2259:10.1016/j.ajhg.2016.10.012 1832: 1549:, as well as responses to 455: 206: 7954: 7934:Evolutionary anthropology 7319: 7126: 6630:10.1038/s41586-020-1929-1 6141:10.1016/j.cub.2021.07.022 4801:10.1142/S0219720018400115 4331:10.1016/j.cub.2017.09.030 4213:21.11116/0000-0007-11C2-A 4086:10.1016/j.gde.2006.09.006 3175:10.1016/j.cub.2023.09.066 2997:10.1101/2024.02.15.580575 2571:10.1007/s11692-015-9348-1 2429:10.1016/j.cub.2016.03.037 1705:shape, a high mandibular 1426:primary biliary cirrhosis 1391: 475: 464: 5604:22 February 2009 at the 5401:Steensby, H. P. (1907). 5346:Annales de Paléontologie 4139:Quaternary International 1985:Archaic African hominins 1948:Changes in modern humans 1820:skeletons from Romania. 1794:Out of Africa hypothesis 1442:diabetes mellitus type 2 1436:size, smoking behavior, 452:Changes in modern humans 7840:Origin of modern humans 6783:7 December 2020 at the 6521:10.1073/pnas.1109300108 6363:10.1126/science.aad2879 6202:10.1073/pnas.1221359110 6000:10.1073/pnas.1108181108 5941:10.1126/science.1244869 5828:10.1126/science.1211177 5587:The Neanderthal within. 5403:"Racestudier i Danmark" 5390:(subscription required) 5224:10.1073/pnas.0702214104 5165:10.1073/pnas.2035108100 5067:10.1073/pnas.0608443103 5003:10.1073/pnas.96.13.7604 4825:Annals of Human Biology 4632:10.1126/science.1209202 4504:10.1073/pnas.0606966103 4204:10.1126/science.abb6460 4122:(1): e1. Archived from 3243:10.1126/science.1224344 3087:10.1126/science.1245938 2852:10.1126/science.aao1887 2666:10.1126/science.1188021 2513:10.1126/science.aay5012 2112:10.3390/biology11081163 2040:artificial intelligence 1864:It has been shown that 1860:Proportion of admixture 1783:Race studies in Denmark 1379:P a r a n t h r o p u s 1238:Dispersal beyond Africa 214:Proportion of admixture 6890:10.1126/sciadv.aay5483 6826:10.1126/sciadv.aax5097 6745:10.1126/sciadv.aax5097 5543:(6941): 692–693, 695. 3117:Barras, Colin (2017). 2074:Neanderthal extinction 1856: 1729: 1663:linkage disequilibrium 1593:adjacent to the right 510:−10 — 458:Recent human evolution 343: 230: 37: 7848:Recent African origin 7086:Last common ancestors 6952:Nature Communications 5692:. 13 September 2006. 5442:"The Origin of races" 5197:Trinkaus, E. (2007). 4868:10.1093/molbev/mss117 4448:10.1093/molbev/mst260 2991:: 2024.02.15.580575. 1912:from Denisovans. The 1842: 1833:Further information: 1723: 1599:primary visual cortex 600:−1 — 590:−2 — 580:−3 — 570:−4 — 560:−5 — 550:−6 — 540:−7 — 530:−8 — 520:−9 — 456:Further information: 443:Tianyuan modern human 334: 221: 35: 7863:Behavioral modernity 7853:Multiregional origin 7633:archaic Homo sapiens 7628:Homo heidelbergensis 7573:Red Deer Cave people 5411:Geographical Journal 4390:10.3390/genes9070358 4317:(20): 3202–3208.e9. 3169:(22): 4905–4916.e5. 2598:(14 December 2023). 2551:Evolutionary Biology 1939:Reduced contribution 1918:Australo-Melanesians 1607:orbitofrontal cortex 1595:intraparietal sulcus 1571:intraparietal sulcus 1543:rheumatoid arthritis 1194:Earliest stone tools 432:background selection 407:Reduced contribution 327:Distance to lineages 209:Neanderthal genetics 43:occurred during the 7500:H. neanderthalensis 7420:H. e. tautavelensis 6965:2019NatCo..10..246M 6882:2020SciA....6.5483R 6818:2020SciA....6.5097D 6736:2020SciA....6.5097D 6679:2022Natur.603..290L 6622:2020Natur.577..665L 6512:2011PNAS..10815123H 6506:(37): 15123–15128. 6265:10.1038/nature13408 6257:2014Natur.512..194H 6193:2013PNAS..110.2223F 6133:2021CBio...31E4219L 5991:2011PNAS..10818301S 5985:(45): 18301–18306. 5933:2013Sci...342..321C 5820:2011Sci...334...94R 5751:10.1038/nature09710 5743:2010Natur.468.1053R 5737:(7327): 1053–1060. 5627:TalkOrigins Archive 5623:Fossil Hominids FAQ 5549:2003Natur.423..692S 5377:1928SciAm.139..112S 5365:Scientific American 5313:Huxley, T. (1890). 5284:10.1038/nature14134 5276:2015Natur.520..216H 5215:2007PNAS..104.7367T 5156:2003PNAS..10011231T 5150:(20): 11231–11236. 5058:2006PNAS..10317196S 5052:(46): 17196–17201. 4994:1999PNAS...96.7604D 4932:2012arXiv1208.2238S 4748:2017NatSR...7.6308G 4624:2011Sci...334...89A 4554:2010PLoSO...510648L 4495:2006PNAS..10318178E 4489:(48): 18178–18183. 4323:2017CBio...27E3202Y 4264:2012PLoSO...747076N 4196:2020Sci...369.1653P 4190:(6511): 1653–1656. 4151:2013QuInt.295..126W 4126:on 6 December 2019. 3937:10.1038/nature16544 3929:2016Natur.530..429K 3782:2012PLoSO...747765S 3714:2012PLoSO...747765S 3651:2012PLoSO...747765S 3588:2012PLoSO...747765S 3441:10.1038/nature21347 3433:2017Natur.541..302N 3375:10.1038/nature12961 3367:2014Natur.507..354S 3235:2012Sci...338..222M 3078:2014Sci...343.1017V 3072:(6174): 1017–1021. 2950:(31 January 2020). 2844:2017Sci...358..655P 2732:10.1038/nature12886 2724:2014Natur.505...43P 2658:2010Sci...328..710G 2563:2016EvBio..43....1A 2421:2016CBio...26.1241S 2204:2020Sci...367..497P 2050:, in the genome of 1806:skeleton of a child 1779:Hans Peder Steensby 1695:superciliary arches 1575:cortical complexity 1573:and an increase in 380:early modern humans 287:purifying selection 263:Introgressed genome 178:In Africa, archaic 163:populations of the 7687:Self-domestication 7478:H. heidelbergensis 7427:H. e. yuanmouensis 7392:H. e. lantianensis 7119:Australopithecines 4837:10.17863/CAM.39003 4736:Scientific Reports 2957:The New York Times 2910:(4): 677–687.e16. 2605:The New York Times 2507:(6484): eaay5012. 1857: 1738:mandibular foramen 1730: 1707:coronoid processus 995:H. heidelbergensis 344: 231: 120:, intermediate in 91:Sub-Saharan Africa 45:Middle Paleolithic 38: 7998: 7997: 7939:Paleoanthropology 7881: 7880: 7858:Archaic admixture 7736:Stoned ape theory 7672:Endurance running 7589: 7588: 7585: 7584: 7581: 7580: 7436: 7435: 7399:H. e. nankinensis 7355:H. tsaichangensis 7291: 7290: 6673:(7900): 290–296. 6616:(7792): 665–670. 6357:(6262): 820–822. 6251:(7513): 194–197. 6127:(19): 4219–4230. 5927:(6156): 321–323. 5689:john hawks weblog 5270:(7546): 216–219. 5209:(18): 7367–7372. 4988:(13): 7604–7609. 4862:(10): 2987–2995. 3923:(7591): 429–433. 3427:(7637): 302–310. 3361:(7492): 354–357. 3229:(6104): 222–226. 2838:(6363): 655–658. 2652:(5979): 710–722. 1724:The modern human 1687:Peștera Muierilor 1404: 1403: 1396:million years ago 1355: 1354: 1333: 1332: 1311: 1310: 1303:Earliest rock art 1289: 1288: 1265: 1264: 1258:Earliest language 1245: 1244: 1225: 1224: 1201: 1200: 1181: 1180: 1171:Earliest sign of 1158: 1157: 1148:Earliest sign of 1135: 1134: 1115: 1114: 1095: 1094: 1075: 1074: 718:Ou. macedoniensis 468:Hominine timeline 392:mitochondrial DNA 304:F4 ancestry ratio 292:Western Eurasians 270:Altai Neanderthal 235:genome sequencing 49:Upper Paleolithic 16:(Redirected from 8033: 8021:Middle Stone Age 7986: 7974: 7973: 7962: 7961: 7898:Human prehistory 7873:Recent evolution 7868:Early migrations 7810:Thermoregulation 7711:Expensive tissue 7682:Sexual selection 7652: 7524: 7406:H. e. pekinensis 7317: 7310: 7225:A. bahrelghazali 7194:Australopithecus 7124: 7094:Chimpanzee–human 7082: 7056: 7049: 7042: 7033: 7026: 7025: 7023: 7021: 7010:ScienceAlert.com 7001: 6995: 6994: 6984: 6942: 6936: 6935: 6933: 6921: 6912: 6911: 6901: 6870:Science Advances 6861: 6848: 6847: 6837: 6806:Science Advances 6797: 6791: 6775: 6769: 6767: 6757: 6747: 6724:Science Advances 6715: 6709: 6708: 6698: 6658: 6652: 6651: 6641: 6601: 6595: 6594: 6584: 6574: 6550: 6544: 6543: 6533: 6523: 6491: 6485: 6484: 6448: 6442: 6441: 6431: 6414:(1): 59–71.e21. 6399: 6393: 6392: 6374: 6345: 6339: 6338: 6328: 6296: 6287: 6286: 6276: 6236: 6225: 6224: 6214: 6204: 6187:(6): 2223–2227. 6172: 6163: 6162: 6152: 6112: 6106: 6105: 6095: 6063: 6054: 6053: 6051: 6049: 6029: 6023: 6022: 6012: 6002: 5970: 5961: 5960: 5916: 5910: 5909: 5899: 5867: 5850: 5849: 5839: 5799: 5793: 5792: 5790: 5788: 5770: 5728: 5719: 5706: 5705: 5703: 5701: 5680: 5674: 5673: 5671: 5669: 5649: 5643: 5642: 5640: 5638: 5614: 5608: 5583: 5577: 5576: 5530: 5524: 5522: 5512: 5477: 5471: 5470: 5444: 5436: 5430: 5429: 5427: 5425: 5419: 5408: 5398: 5392: 5391: 5388: 5360: 5354: 5353: 5341: 5335: 5334: 5332: 5330: 5310: 5304: 5303: 5258: 5247: 5246: 5236: 5226: 5194: 5188: 5187: 5177: 5167: 5135: 5124: 5123: 5121: 5119: 5104: 5098: 5097: 5087: 5069: 5037: 5026: 5025: 5015: 5005: 4973: 4964: 4963: 4953: 4943: 4925: 4916:(10): e1002947. 4901: 4890: 4889: 4879: 4847: 4841: 4840: 4819: 4813: 4812: 4784: 4778: 4777: 4767: 4727: 4710: 4709: 4699: 4667: 4654: 4653: 4643: 4603: 4586: 4585: 4575: 4565: 4533: 4527: 4526: 4516: 4506: 4474: 4461: 4460: 4450: 4426: 4413: 4412: 4402: 4392: 4368: 4353: 4352: 4342: 4302: 4296: 4295: 4285: 4275: 4243: 4234: 4233: 4215: 4181: 4172: 4163: 4162: 4134: 4128: 4127: 4107: 4098: 4097: 4069: 4063: 4062: 4052: 4042: 4018: 4012: 4011: 3993: 3965: 3959: 3958: 3948: 3908: 3897: 3896: 3894: 3892: 3876: 3870: 3869: 3856: 3824: 3818: 3817: 3803: 3793: 3761: 3755: 3754: 3750: 3735: 3725: 3693: 3687: 3686: 3672: 3662: 3630: 3624: 3623: 3609: 3599: 3567: 3561: 3560: 3550: 3518: 3512: 3511: 3501: 3469: 3463: 3462: 3452: 3412: 3397: 3396: 3386: 3346: 3321: 3320: 3310: 3278: 3265: 3264: 3254: 3220: 3211: 3194: 3193: 3191: 3189: 3153: 3147: 3146: 3136: 3134: 3114: 3108: 3107: 3089: 3057: 3038: 3037: 3025: 3019: 3018: 3008: 2980: 2974: 2973: 2971: 2969: 2944: 2938: 2937: 2919: 2895: 2874: 2873: 2863: 2823: 2806: 2805: 2795: 2778:(4): 1241–1251. 2763: 2754: 2753: 2743: 2703: 2688: 2687: 2677: 2637: 2622: 2621: 2619: 2617: 2592: 2583: 2582: 2542: 2536: 2534: 2524: 2490: 2481: 2480: 2478: 2476: 2457: 2451: 2450: 2440: 2415:(9): 1241–1247. 2396: 2379: 2378: 2368: 2358: 2334: 2295: 2294: 2280: 2270: 2253:(6): 1316–1324. 2238: 2232: 2231: 2187: 2176: 2175: 2173: 2171: 2159:Business Insider 2149: 2143: 2142: 2132: 2114: 2090: 2036:genetics studies 1703:mandibular notch 1653:allele frequency 1567:temporal locales 1539:eating disorders 1415:Genes affecting 1384: 1382: 1381: 1367: 1365: 1349: 1340: 1335: 1327: 1325:Earliest clothes 1318: 1313: 1305: 1296: 1291: 1272: 1267: 1252: 1247: 1232: 1227: 1214:Earliest sign of 1208: 1203: 1188: 1183: 1173:Australopithecus 1165: 1160: 1142: 1137: 1128:Earliest bipedal 1122: 1117: 1108:Chimpanzee split 1102: 1097: 1082: 1077: 1062: 1057: 1043: 1042: 1028: 1027: 1011: 997: 983: 955: 942: 922: 909: 881: 879:Australopithecus 868: 853: 836: 823: 799: 786: 773: 760: 747: 734: 722: 703: 690: 677: 665: 651: 638: 625: 623: 611: 606: 601: 596: 591: 586: 581: 576: 571: 566: 561: 556: 551: 546: 541: 536: 531: 526: 521: 516: 511: 499: 492: 485: 479: 469: 462: 142:Native Americans 126:Southeast Asians 21: 8041: 8040: 8036: 8035: 8034: 8032: 8031: 8030: 8001: 8000: 7999: 7994: 7950: 7907: 7893:Human evolution 7877: 7834: 7778: 7772: 7751:Cooperative eye 7696:Specific models 7691: 7643: 7622:Homo antecessor 7577: 7513: 7507:H. rhodesiensis 7471:H. floresiensis 7432: 7413:H. e. soloensis 7385:H. e. georgicus 7361: 7325:H. gautengensis 7300: 7298: 7287: 7251: 7187: 7158: 7113: 7104:Orangutan–human 7073: 7065: 7063:Human evolution 7060: 7030: 7029: 7019: 7017: 7003: 7002: 6998: 6944: 6943: 6939: 6923: 6922: 6915: 6876:(8): eaay5483. 6863: 6862: 6851: 6812:(7): eaax5097. 6799: 6798: 6794: 6785:Wayback Machine 6776: 6772: 6730:(7): eaax5097. 6717: 6716: 6712: 6660: 6659: 6655: 6603: 6602: 6598: 6552: 6551: 6547: 6493: 6492: 6488: 6465:10.1038/nrg3295 6459:(10): 745–753. 6453:Nat. Rev. Genet 6450: 6449: 6445: 6401: 6400: 6396: 6347: 6346: 6342: 6298: 6297: 6290: 6238: 6237: 6228: 6174: 6173: 6166: 6121:Current Biology 6114: 6113: 6109: 6078:(1): 53–61.e9. 6065: 6064: 6057: 6047: 6045: 6031: 6030: 6026: 5972: 5971: 5964: 5918: 5917: 5913: 5869: 5868: 5853: 5814:(6052): 94–98. 5801: 5800: 5796: 5786: 5784: 5726: 5721: 5720: 5709: 5699: 5697: 5682: 5681: 5677: 5667: 5665: 5651: 5650: 5646: 5636: 5634: 5616: 5615: 5611: 5606:Wayback Machine 5584: 5580: 5557:10.1038/423692a 5532: 5531: 5527: 5480: 5478: 5474: 5438: 5437: 5433: 5423: 5421: 5417: 5406: 5400: 5399: 5395: 5389: 5362: 5361: 5357: 5343: 5342: 5338: 5328: 5326: 5312: 5311: 5307: 5260: 5259: 5250: 5196: 5195: 5191: 5137: 5136: 5127: 5117: 5115: 5106: 5105: 5101: 5039: 5038: 5029: 4975: 4974: 4967: 4903: 4902: 4893: 4849: 4848: 4844: 4821: 4820: 4816: 4786: 4785: 4781: 4729: 4728: 4713: 4669: 4668: 4657: 4618:(6052): 89–94. 4605: 4604: 4589: 4535: 4534: 4530: 4476: 4475: 4464: 4428: 4427: 4416: 4370: 4369: 4356: 4311:Current Biology 4304: 4303: 4299: 4245: 4244: 4237: 4179: 4174: 4173: 4166: 4136: 4135: 4131: 4109: 4108: 4101: 4071: 4070: 4066: 4020: 4019: 4015: 3967: 3966: 3962: 3910: 3909: 3900: 3890: 3888: 3878: 3877: 3873: 3826: 3825: 3821: 3763: 3762: 3758: 3748: 3695: 3694: 3690: 3632: 3631: 3627: 3569: 3568: 3564: 3520: 3519: 3515: 3471: 3470: 3466: 3414: 3413: 3400: 3348: 3347: 3324: 3280: 3279: 3268: 3218: 3213: 3212: 3197: 3187: 3185: 3163:Current Biology 3155: 3154: 3150: 3132: 3130: 3116: 3115: 3111: 3059: 3058: 3041: 3027: 3026: 3022: 2982: 2981: 2977: 2967: 2965: 2946: 2945: 2941: 2897: 2896: 2877: 2825: 2824: 2809: 2765: 2764: 2757: 2718:(7481): 43–49. 2705: 2704: 2691: 2639: 2638: 2625: 2615: 2613: 2594: 2593: 2586: 2544: 2543: 2539: 2492: 2491: 2484: 2474: 2472: 2459: 2458: 2454: 2409:Current Biology 2398: 2397: 2382: 2349:(5): e1007349. 2336: 2335: 2298: 2240: 2239: 2235: 2189: 2188: 2179: 2169: 2167: 2151: 2150: 2146: 2092: 2091: 2087: 2082: 2060: 2032: 2030:Related studies 2011: 1987: 1950: 1941: 1862: 1837: 1831: 1826: 1818:Peștera Muierii 1767: 1734:Peștera cu Oase 1676: 1644: 1591:parietal region 1535:LDL cholesterol 1466:coalescence age 1430:Crohn's disease 1400: 1399: 1387: 1386: 1385: 1377: 1376: 1374: 1370: 1369: 1368: 1363:H o m i n i d s 1361: 1359: 1351: 1350: 1345: 1338: 1329: 1328: 1323: 1316: 1307: 1306: 1301: 1294: 1285: 1284: 1270: 1261: 1260: 1250: 1241: 1240: 1230: 1221: 1220: 1215: 1206: 1197: 1196: 1186: 1177: 1176: 1163: 1154: 1153: 1140: 1131: 1130: 1120: 1111: 1110: 1100: 1091: 1090: 1080: 1071: 1070: 1060: 1053: 1052: 1051: 1046: 1045: 1044: 1038: 1037: 1033: 1031: 1030: 1029: 1023: 1022: 1018: 1016: 1015: 1014: 1007: 1001: 1000: 999: 993: 988: 986: 985: 984: 975: 967: 959: 958: 951: 945: 944: 943: 934: 926: 925: 918: 912: 911: 910: 901: 893: 885: 884: 877: 871: 870: 869: 860: 856: 855: 854: 845: 841: 840: 839: 832: 826: 825: 824: 815: 807: 803: 802: 801: 795: 790: 789: 788: 782: 777: 776: 775: 769: 764: 763: 762: 756: 751: 750: 749: 743: 738: 737: 736: 732:Chororapithecus 730: 725: 724: 723: 714: 706: 705: 699: 694: 693: 692: 688:Samburupithecus 686: 681: 680: 679: 673: 668: 667: 666: 659: 655: 654: 653: 647: 642: 641: 640: 634: 629: 628: 627: 621: 619: 612: 609: 607: 604: 602: 599: 597: 594: 592: 589: 587: 584: 582: 579: 577: 574: 572: 569: 567: 564: 562: 559: 557: 554: 552: 549: 547: 544: 542: 539: 537: 534: 532: 529: 527: 524: 522: 519: 517: 514: 512: 509: 503: 477: 471: 467: 460: 454: 409: 388: 329: 278: 265: 216: 211: 205: 200: 124:, and lower in 28: 23: 22: 15: 12: 11: 5: 8039: 8037: 8029: 8028: 8023: 8018: 8013: 8003: 8002: 7996: 7995: 7993: 7992: 7980: 7968: 7955: 7952: 7951: 7949: 7948: 7943: 7942: 7941: 7931: 7926: 7921: 7915: 7913: 7909: 7908: 7906: 7905: 7903:Human timeline 7900: 7895: 7889: 7887: 7883: 7882: 7879: 7878: 7876: 7875: 7870: 7865: 7860: 7855: 7850: 7844: 7842: 7836: 7835: 7833: 7832: 7827: 7822: 7817: 7812: 7807: 7802: 7797: 7792: 7787: 7781: 7779: 7774: 7773: 7771: 7770: 7769: 7768: 7763: 7755: 7754: 7753: 7748: 7740: 7739: 7738: 7733: 7728: 7726:Drunken monkey 7720: 7719: 7718: 7713: 7708: 7699: 7697: 7693: 7692: 7690: 7689: 7684: 7679: 7674: 7669: 7664: 7658: 7656: 7655:General models 7649: 7645: 7644: 7642: 7641: 7599: 7597: 7591: 7590: 7587: 7586: 7583: 7582: 7579: 7578: 7576: 7575: 7570: 7565: 7560: 7555: 7548: 7543: 7534: 7532: 7521: 7515: 7514: 7512: 7511: 7503: 7496: 7489: 7481: 7474: 7467: 7459: 7454: 7446: 7444: 7442:Archaic humans 7438: 7437: 7434: 7433: 7431: 7430: 7423: 7416: 7409: 7402: 7395: 7388: 7381: 7373: 7371: 7363: 7362: 7360: 7359: 7351: 7347:H. rudolfensis 7343: 7336: 7329: 7320: 7314: 7307: 7293: 7292: 7289: 7288: 7286: 7285: 7278: 7271: 7268:P. aethiopicus 7263: 7261: 7253: 7252: 7250: 7249: 7242: 7235: 7228: 7221: 7214: 7207: 7199: 7197: 7189: 7188: 7186: 7185: 7178: 7170: 7168: 7160: 7159: 7157: 7156: 7149: 7146:Sahelanthropus 7142: 7135: 7132:Nakalipithecus 7127: 7121: 7115: 7114: 7112: 7111: 7106: 7101: 7096: 7090: 7088: 7079: 7067: 7066: 7061: 7059: 7058: 7051: 7044: 7036: 7028: 7027: 6996: 6937: 6913: 6849: 6792: 6770: 6710: 6653: 6596: 6545: 6486: 6443: 6394: 6340: 6311:(3): 457–469. 6288: 6226: 6164: 6107: 6055: 6024: 5962: 5911: 5882:(4): 516–528. 5851: 5794: 5707: 5675: 5644: 5609: 5578: 5525: 5501:10.1086/505436 5495:(2): 230–237. 5472: 5431: 5393: 5371:(2): 112–115. 5355: 5336: 5305: 5248: 5189: 5125: 5099: 5027: 4965: 4891: 4842: 4814: 4795:(2): 1840011. 4779: 4711: 4682:(5): 916–927. 4655: 4587: 4528: 4462: 4441:(3): 683–695. 4414: 4354: 4297: 4258:(10): e47076. 4235: 4164: 4129: 4099: 4080:(6): 606–610. 4064: 4033:(3): 313–317. 4013: 3960: 3898: 3871: 3839:(1): 199–209. 3819: 3776:(10): e47765. 3756: 3708:(10): e47765. 3688: 3645:(10): e47765. 3625: 3582:(10): e47765. 3562: 3533:(3): 454–461. 3513: 3484:(3): 448–453. 3464: 3398: 3322: 3293:(1): 199–209. 3266: 3195: 3148: 3109: 3039: 3020: 2975: 2939: 2875: 2807: 2755: 2689: 2623: 2584: 2537: 2482: 2452: 2380: 2296: 2247:Am J Hum Genet 2233: 2177: 2144: 2084: 2083: 2081: 2078: 2077: 2076: 2071: 2066: 2059: 2056: 2031: 2028: 2010: 2007: 1986: 1983: 1949: 1946: 1940: 1937: 1861: 1858: 1830: 1827: 1825: 1822: 1766: 1763: 1697:, a prominent 1675: 1672: 1646:Although less 1643: 1640: 1577:for the early 1563:parietal bones 1487:cis-regulatory 1438:interleukin 18 1402: 1401: 1393: 1392: 1389: 1388: 1373: 1372: 1371: 1358: 1357: 1356: 1353: 1352: 1344: 1343: 1341: 1331: 1330: 1322: 1321: 1319: 1309: 1308: 1300: 1299: 1297: 1287: 1286: 1276: 1275: 1273: 1263: 1262: 1256: 1255: 1253: 1243: 1242: 1236: 1235: 1233: 1223: 1222: 1212: 1211: 1209: 1199: 1198: 1192: 1191: 1189: 1179: 1178: 1169: 1168: 1166: 1156: 1155: 1146: 1145: 1143: 1133: 1132: 1126: 1125: 1123: 1113: 1112: 1106: 1105: 1103: 1093: 1092: 1086: 1085: 1083: 1073: 1072: 1066: 1065: 1063: 1054: 1049: 1048: 1047: 1036: 1035: 1034: 1032: 1021: 1020: 1019: 1017: 1004: 1003: 1002: 991: 990: 989: 987: 948: 947: 946: 930:H. rudolfensis 915: 914: 913: 874: 873: 872: 859: 858: 857: 844: 843: 842: 829: 828: 827: 806: 805: 804: 793: 792: 791: 784:Graecopithecus 780: 779: 778: 771:Sahelanthropus 767: 766: 765: 754: 753: 752: 741: 740: 739: 728: 727: 726: 701:Ouranopithecus 697: 696: 695: 684: 683: 682: 675:Nakalipithecus 671: 670: 669: 658: 657: 656: 645: 644: 643: 632: 631: 630: 617: 616: 615: 613: 610:0 — 608: 603: 598: 593: 588: 583: 578: 573: 568: 563: 558: 553: 548: 543: 538: 533: 528: 523: 518: 513: 508: 505: 504: 502: 501: 494: 487: 476: 473: 472: 465: 453: 450: 408: 405: 387: 384: 374:of the Altai, 354:) than to the 352:North Caucasus 328: 325: 277: 274: 264: 261: 215: 212: 207:Main article: 204: 201: 199: 196: 111:West Eurasians 103:Horn of Africa 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 8038: 8027: 8024: 8022: 8019: 8017: 8014: 8012: 8011:Human hybrids 8009: 8008: 8006: 7991: 7990: 7985: 7981: 7979: 7978: 7969: 7967: 7966: 7957: 7956: 7953: 7947: 7944: 7940: 7937: 7936: 7935: 7932: 7930: 7927: 7925: 7922: 7920: 7917: 7916: 7914: 7910: 7904: 7901: 7899: 7896: 7894: 7891: 7890: 7888: 7884: 7874: 7871: 7869: 7866: 7864: 7861: 7859: 7856: 7854: 7851: 7849: 7846: 7845: 7843: 7841: 7837: 7831: 7828: 7826: 7823: 7821: 7818: 7816: 7813: 7811: 7808: 7806: 7803: 7801: 7798: 7796: 7793: 7791: 7788: 7786: 7783: 7782: 7780: 7775: 7767: 7764: 7762: 7759: 7758: 7757:Life history 7756: 7752: 7749: 7747: 7744: 7743: 7741: 7737: 7734: 7732: 7729: 7727: 7724: 7723: 7721: 7717: 7714: 7712: 7709: 7707: 7704: 7703: 7701: 7700: 7698: 7694: 7688: 7685: 7683: 7680: 7678: 7675: 7673: 7670: 7668: 7665: 7663: 7660: 7659: 7657: 7653: 7650: 7646: 7640: 7639: 7634: 7630: 7629: 7624: 7623: 7618: 7617: 7612: 7611: 7610:Homo ergaster 7606: 7605: 7601: 7600: 7598: 7596: 7592: 7574: 7571: 7569: 7566: 7564: 7561: 7559: 7556: 7554: 7553: 7549: 7547: 7544: 7542: 7540: 7539:H. s. sapiens 7536: 7535: 7533: 7531: 7530: 7525: 7522: 7520: 7519:Modern humans 7516: 7509: 7508: 7504: 7502: 7501: 7497: 7495: 7494: 7493:H. luzonensis 7490: 7487: 7486: 7482: 7480: 7479: 7475: 7473: 7472: 7468: 7465: 7464: 7460: 7458: 7455: 7453: 7452: 7451:H. antecessor 7448: 7447: 7445: 7443: 7439: 7429: 7428: 7424: 7422: 7421: 7417: 7415: 7414: 7410: 7408: 7407: 7403: 7401: 7400: 7396: 7394: 7393: 7389: 7387: 7386: 7382: 7380: 7379: 7378:H. e. erectus 7375: 7374: 7372: 7370: 7369: 7364: 7357: 7356: 7352: 7349: 7348: 7344: 7342: 7341: 7337: 7335: 7334: 7330: 7327: 7326: 7322: 7321: 7318: 7315: 7311: 7308: 7306: 7304: 7294: 7284: 7283: 7279: 7277: 7276: 7272: 7270: 7269: 7265: 7264: 7262: 7260: 7259: 7254: 7248: 7247: 7243: 7241: 7240: 7236: 7234: 7233: 7232:A. deyiremeda 7229: 7227: 7226: 7222: 7220: 7219: 7215: 7213: 7212: 7208: 7206: 7205: 7201: 7200: 7198: 7196: 7195: 7190: 7184: 7183: 7179: 7177: 7176: 7172: 7171: 7169: 7167: 7166: 7161: 7155: 7154: 7153:Kenyanthropus 7150: 7148: 7147: 7143: 7141: 7140: 7136: 7134: 7133: 7129: 7128: 7125: 7122: 7120: 7116: 7110: 7107: 7105: 7102: 7100: 7099:Gorilla–human 7097: 7095: 7092: 7091: 7089: 7087: 7083: 7080: 7077: 7072: 7068: 7064: 7057: 7052: 7050: 7045: 7043: 7038: 7037: 7034: 7015: 7011: 7007: 7000: 6997: 6992: 6988: 6983: 6978: 6974: 6970: 6966: 6962: 6958: 6954: 6953: 6948: 6941: 6938: 6932: 6927: 6920: 6918: 6914: 6909: 6905: 6900: 6895: 6891: 6887: 6883: 6879: 6875: 6871: 6867: 6860: 6858: 6856: 6854: 6850: 6845: 6841: 6836: 6831: 6827: 6823: 6819: 6815: 6811: 6807: 6803: 6796: 6793: 6788: 6786: 6782: 6779: 6774: 6771: 6765: 6761: 6756: 6751: 6746: 6741: 6737: 6733: 6729: 6725: 6721: 6714: 6711: 6706: 6702: 6697: 6692: 6688: 6684: 6680: 6676: 6672: 6668: 6664: 6657: 6654: 6649: 6645: 6640: 6635: 6631: 6627: 6623: 6619: 6615: 6611: 6607: 6600: 6597: 6592: 6588: 6583: 6578: 6573: 6568: 6564: 6560: 6559:PLOS Genetics 6556: 6549: 6546: 6541: 6537: 6532: 6527: 6522: 6517: 6513: 6509: 6505: 6501: 6497: 6490: 6487: 6482: 6478: 6474: 6470: 6466: 6462: 6458: 6454: 6447: 6444: 6439: 6435: 6430: 6425: 6421: 6417: 6413: 6409: 6405: 6398: 6395: 6390: 6386: 6382: 6378: 6373: 6368: 6364: 6360: 6356: 6352: 6344: 6341: 6336: 6332: 6327: 6322: 6318: 6314: 6310: 6306: 6302: 6295: 6293: 6289: 6284: 6280: 6275: 6270: 6266: 6262: 6258: 6254: 6250: 6246: 6242: 6235: 6233: 6231: 6227: 6222: 6218: 6213: 6208: 6203: 6198: 6194: 6190: 6186: 6182: 6178: 6171: 6169: 6165: 6160: 6156: 6151: 6146: 6142: 6138: 6134: 6130: 6126: 6122: 6118: 6111: 6108: 6103: 6099: 6094: 6089: 6085: 6081: 6077: 6073: 6069: 6062: 6060: 6056: 6043: 6039: 6035: 6028: 6025: 6020: 6016: 6011: 6006: 6001: 5996: 5992: 5988: 5984: 5980: 5976: 5969: 5967: 5963: 5958: 5954: 5950: 5946: 5942: 5938: 5934: 5930: 5926: 5922: 5915: 5912: 5907: 5903: 5898: 5893: 5889: 5885: 5881: 5877: 5873: 5866: 5864: 5862: 5860: 5858: 5856: 5852: 5847: 5843: 5838: 5833: 5829: 5825: 5821: 5817: 5813: 5809: 5805: 5798: 5795: 5782: 5778: 5774: 5769: 5764: 5760: 5756: 5752: 5748: 5744: 5740: 5736: 5732: 5725: 5718: 5716: 5714: 5712: 5708: 5695: 5691: 5690: 5685: 5679: 5676: 5663: 5659: 5655: 5648: 5645: 5632: 5628: 5624: 5620: 5613: 5610: 5607: 5603: 5600: 5596: 5592: 5591:New Scientist 5588: 5582: 5579: 5574: 5570: 5566: 5562: 5558: 5554: 5550: 5546: 5542: 5538: 5537: 5529: 5526: 5520: 5516: 5511: 5506: 5502: 5498: 5494: 5490: 5489: 5484: 5476: 5473: 5468: 5464: 5460: 5456: 5452: 5448: 5443: 5435: 5432: 5416: 5412: 5404: 5397: 5394: 5386: 5382: 5378: 5374: 5370: 5366: 5359: 5356: 5351: 5348:(in French). 5347: 5340: 5337: 5324: 5320: 5316: 5309: 5306: 5301: 5297: 5293: 5289: 5285: 5281: 5277: 5273: 5269: 5265: 5257: 5255: 5253: 5249: 5244: 5240: 5235: 5230: 5225: 5220: 5216: 5212: 5208: 5204: 5200: 5193: 5190: 5185: 5181: 5176: 5171: 5166: 5161: 5157: 5153: 5149: 5145: 5141: 5134: 5132: 5130: 5126: 5113: 5109: 5103: 5100: 5095: 5091: 5086: 5081: 5077: 5073: 5068: 5063: 5059: 5055: 5051: 5047: 5043: 5036: 5034: 5032: 5028: 5023: 5019: 5014: 5009: 5004: 4999: 4995: 4991: 4987: 4983: 4979: 4972: 4970: 4966: 4961: 4957: 4952: 4947: 4942: 4937: 4933: 4929: 4924: 4919: 4915: 4911: 4910:PLOS Genetics 4907: 4900: 4898: 4896: 4892: 4887: 4883: 4878: 4873: 4869: 4865: 4861: 4857: 4853: 4846: 4843: 4838: 4834: 4831:(2): 99–108. 4830: 4826: 4818: 4815: 4810: 4806: 4802: 4798: 4794: 4790: 4783: 4780: 4775: 4771: 4766: 4761: 4757: 4753: 4749: 4745: 4741: 4737: 4733: 4726: 4724: 4722: 4720: 4718: 4716: 4712: 4707: 4703: 4698: 4693: 4689: 4685: 4681: 4677: 4673: 4666: 4664: 4662: 4660: 4656: 4651: 4647: 4642: 4637: 4633: 4629: 4625: 4621: 4617: 4613: 4609: 4602: 4600: 4598: 4596: 4594: 4592: 4588: 4583: 4579: 4574: 4569: 4564: 4559: 4555: 4551: 4548:(5): e10648. 4547: 4543: 4539: 4532: 4529: 4524: 4520: 4515: 4510: 4505: 4500: 4496: 4492: 4488: 4484: 4480: 4473: 4471: 4469: 4467: 4463: 4458: 4454: 4449: 4444: 4440: 4436: 4432: 4425: 4423: 4421: 4419: 4415: 4410: 4406: 4401: 4396: 4391: 4386: 4382: 4378: 4374: 4367: 4365: 4363: 4361: 4359: 4355: 4350: 4346: 4341: 4336: 4332: 4328: 4324: 4320: 4316: 4312: 4308: 4301: 4298: 4293: 4289: 4284: 4279: 4274: 4269: 4265: 4261: 4257: 4253: 4249: 4242: 4240: 4236: 4231: 4227: 4223: 4219: 4214: 4209: 4205: 4201: 4197: 4193: 4189: 4185: 4178: 4171: 4169: 4165: 4160: 4156: 4152: 4148: 4144: 4140: 4133: 4130: 4125: 4121: 4117: 4113: 4106: 4104: 4100: 4095: 4091: 4087: 4083: 4079: 4075: 4068: 4065: 4060: 4056: 4051: 4046: 4041: 4036: 4032: 4028: 4024: 4017: 4014: 4009: 4005: 4001: 3997: 3992: 3987: 3983: 3979: 3975: 3971: 3964: 3961: 3956: 3952: 3947: 3942: 3938: 3934: 3930: 3926: 3922: 3918: 3914: 3907: 3905: 3903: 3899: 3886: 3882: 3875: 3872: 3868: 3864: 3860: 3855: 3850: 3846: 3842: 3838: 3834: 3830: 3823: 3820: 3816: 3811: 3807: 3802: 3797: 3792: 3787: 3783: 3779: 3775: 3771: 3767: 3760: 3757: 3753: 3743: 3739: 3734: 3729: 3724: 3719: 3715: 3711: 3707: 3703: 3699: 3692: 3689: 3685: 3680: 3676: 3671: 3666: 3661: 3656: 3652: 3648: 3644: 3640: 3636: 3629: 3626: 3622: 3617: 3613: 3608: 3603: 3598: 3593: 3589: 3585: 3581: 3577: 3573: 3566: 3563: 3558: 3554: 3549: 3544: 3540: 3536: 3532: 3528: 3524: 3517: 3514: 3509: 3505: 3500: 3495: 3491: 3487: 3483: 3479: 3475: 3468: 3465: 3460: 3456: 3451: 3446: 3442: 3438: 3434: 3430: 3426: 3422: 3418: 3411: 3409: 3407: 3405: 3403: 3399: 3394: 3390: 3385: 3380: 3376: 3372: 3368: 3364: 3360: 3356: 3352: 3345: 3343: 3341: 3339: 3337: 3335: 3333: 3331: 3329: 3327: 3323: 3318: 3314: 3309: 3304: 3300: 3296: 3292: 3288: 3284: 3277: 3275: 3273: 3271: 3267: 3262: 3258: 3253: 3248: 3244: 3240: 3236: 3232: 3228: 3224: 3217: 3210: 3208: 3206: 3204: 3202: 3200: 3196: 3184: 3180: 3176: 3172: 3168: 3164: 3160: 3152: 3149: 3145: 3143: 3128: 3124: 3123:New Scientist 3120: 3113: 3110: 3105: 3101: 3097: 3093: 3088: 3083: 3079: 3075: 3071: 3067: 3063: 3056: 3054: 3052: 3050: 3048: 3046: 3044: 3040: 3035: 3034:New Scientist 3031: 3024: 3021: 3016: 3012: 3007: 3002: 2998: 2994: 2990: 2986: 2979: 2976: 2963: 2959: 2958: 2953: 2949: 2943: 2940: 2935: 2931: 2927: 2923: 2918: 2913: 2909: 2905: 2901: 2894: 2892: 2890: 2888: 2886: 2884: 2882: 2880: 2876: 2871: 2867: 2862: 2857: 2853: 2849: 2845: 2841: 2837: 2833: 2829: 2822: 2820: 2818: 2816: 2814: 2812: 2808: 2803: 2799: 2794: 2789: 2785: 2781: 2777: 2773: 2769: 2762: 2760: 2756: 2751: 2747: 2742: 2737: 2733: 2729: 2725: 2721: 2717: 2713: 2709: 2702: 2700: 2698: 2696: 2694: 2690: 2685: 2681: 2676: 2671: 2667: 2663: 2659: 2655: 2651: 2647: 2643: 2636: 2634: 2632: 2630: 2628: 2624: 2611: 2607: 2606: 2601: 2597: 2591: 2589: 2585: 2580: 2576: 2572: 2568: 2564: 2560: 2556: 2552: 2548: 2541: 2538: 2532: 2528: 2523: 2518: 2514: 2510: 2506: 2502: 2501: 2496: 2489: 2487: 2483: 2470: 2466: 2462: 2456: 2453: 2448: 2444: 2439: 2434: 2430: 2426: 2422: 2418: 2414: 2410: 2406: 2402: 2395: 2393: 2391: 2389: 2387: 2385: 2381: 2376: 2372: 2367: 2362: 2357: 2352: 2348: 2344: 2343:PLOS Genetics 2340: 2333: 2331: 2329: 2327: 2325: 2323: 2321: 2319: 2317: 2315: 2313: 2311: 2309: 2307: 2305: 2303: 2301: 2297: 2293: 2288: 2284: 2279: 2274: 2269: 2268:11577/3455530 2264: 2260: 2256: 2252: 2248: 2244: 2237: 2234: 2229: 2225: 2221: 2217: 2213: 2209: 2205: 2201: 2198:(6477): 497. 2197: 2193: 2186: 2184: 2182: 2178: 2165: 2161: 2160: 2155: 2148: 2145: 2140: 2136: 2131: 2126: 2122: 2118: 2113: 2108: 2104: 2100: 2096: 2089: 2086: 2079: 2075: 2072: 2070: 2067: 2065: 2064:Interbreeding 2062: 2061: 2057: 2055: 2053: 2052:modern humans 2049: 2045: 2041: 2037: 2029: 2027: 2023: 2019: 2015: 2008: 2006: 2002: 1998: 1994: 1990: 1984: 1982: 1979: 1975: 1971: 1967: 1962: 1958: 1955: 1947: 1945: 1938: 1936: 1934: 1930: 1925: 1921: 1919: 1915: 1914:Ayta Magbukon 1911: 1906: 1902: 1898: 1896: 1890: 1886: 1884: 1878: 1876: 1871: 1867: 1859: 1854: 1850: 1846: 1841: 1836: 1828: 1823: 1821: 1819: 1815: 1811: 1807: 1803: 1799: 1798:Erik Trinkaus 1795: 1790: 1786: 1784: 1780: 1775: 1772: 1771:Thomas Huxley 1764: 1762: 1760: 1756: 1752: 1748: 1746: 1741: 1739: 1735: 1727: 1722: 1718: 1716: 1715:glenoid fossa 1712: 1708: 1704: 1700: 1699:occipital bun 1696: 1692: 1688: 1683: 1681: 1673: 1671: 1669: 1664: 1659: 1654: 1649: 1641: 1639: 1637: 1633: 1629: 1628:modern humans 1626:inherited by 1625: 1620: 1617: 1615: 1610: 1608: 1603: 1600: 1596: 1592: 1588: 1584: 1580: 1579:visual cortex 1576: 1572: 1568: 1565:to bilateral 1564: 1561:and inferior 1560: 1554: 1552: 1551:antipsychotic 1548: 1547:schizophrenia 1544: 1540: 1536: 1531: 1529: 1525: 1521: 1517: 1513: 1509: 1503: 1501: 1500:basal ganglia 1497: 1492: 1488: 1483: 1479: 1477: 1476:Monti Lessini 1472: 1467: 1463: 1459: 1458:microcephalin 1454: 1451: 1445: 1443: 1439: 1435: 1431: 1427: 1423: 1418: 1413: 1410: 1397: 1390: 1383: 1380: 1366: 1364: 1348: 1347:Modern humans 1342: 1337: 1336: 1326: 1320: 1315: 1314: 1304: 1298: 1293: 1292: 1283: 1279: 1278:Earliest fire 1274: 1269: 1268: 1259: 1254: 1249: 1248: 1239: 1234: 1229: 1228: 1219: 1218: 1210: 1205: 1204: 1195: 1190: 1185: 1184: 1175: 1174: 1167: 1162: 1161: 1152: 1151: 1144: 1139: 1138: 1129: 1124: 1119: 1118: 1109: 1104: 1099: 1098: 1089: 1088:Gorilla split 1084: 1079: 1078: 1069: 1064: 1059: 1058: 1055: 1041: 1026: 1013: 1012: 1010: 998: 996: 982: 980: 974: 972: 966: 964: 963:H. antecessor 957: 956: 954: 941: 939: 933: 931: 924: 923: 921: 908: 906: 905:Au. anamensis 900: 898: 897:Au. afarensis 892: 890: 889:Au. africanus 883: 882: 880: 867: 865: 852: 850: 838: 837: 835: 822: 820: 819:O. tugenensis 814: 812: 800: 798: 787: 785: 774: 772: 761: 759: 748: 746: 735: 733: 721: 719: 713: 711: 704: 702: 691: 689: 678: 676: 664: 663: 652: 650: 639: 637: 626: 624: 614: 507: 506: 500: 495: 493: 488: 486: 481: 480: 474: 470: 463: 459: 451: 449: 446: 444: 439: 436: 433: 428: 426: 422: 418: 414: 406: 404: 400: 398: 393: 385: 383: 381: 377: 373: 372:chromosome 21 368: 365: 361: 358:Neanderthal ( 357: 353: 350:Neanderthal ( 349: 341: 337: 333: 326: 324: 321: 317: 313: 309: 305: 299: 295: 293: 288: 284: 275: 273: 271: 262: 260: 258: 253: 248: 245: 240: 236: 228: 224: 220: 213: 210: 202: 197: 195: 193: 188: 184: 181: 176: 174: 170: 166: 162: 158: 154: 150: 145: 143: 139: 135: 131: 127: 123: 119: 114: 112: 108: 104: 100: 96: 92: 87: 85: 84:introgression 81: 80:modern humans 77: 73: 69: 64: 62: 58: 54: 50: 46: 42: 34: 30: 19: 7987: 7975: 7963: 7857: 7830:Gender roles 7825:Intelligence 7638:Homo sapiens 7636: 7632: 7626: 7620: 7616:Homo erectus 7614: 7608: 7604:Homo habilis 7602: 7563:Manot people 7552:H. s. idaltu 7550: 7546:Jebel Irhoud 7538: 7529:Homo sapiens 7527: 7505: 7498: 7491: 7483: 7476: 7469: 7461: 7449: 7425: 7418: 7411: 7404: 7397: 7390: 7383: 7376: 7368:Homo erectus 7366: 7353: 7345: 7338: 7331: 7323: 7313:Proto-humans 7302: 7299:proto-humans 7280: 7273: 7266: 7258:Paranthropus 7256: 7244: 7237: 7230: 7223: 7218:A. anamensis 7216: 7211:A. africanus 7209: 7204:A. afarensis 7202: 7192: 7180: 7173: 7165:Ardipithecus 7163: 7151: 7144: 7137: 7130: 7109:Gibbon–human 7018:. Retrieved 7009: 6999: 6959:(246): 246. 6956: 6950: 6940: 6873: 6869: 6809: 6805: 6795: 6777: 6773: 6727: 6723: 6713: 6670: 6666: 6656: 6613: 6609: 6599: 6562: 6558: 6548: 6503: 6499: 6489: 6456: 6452: 6446: 6411: 6407: 6397: 6372:2318/1661894 6354: 6350: 6343: 6308: 6304: 6248: 6244: 6184: 6180: 6124: 6120: 6110: 6075: 6071: 6046:. Retrieved 6037: 6027: 5982: 5978: 5924: 5920: 5914: 5879: 5875: 5811: 5807: 5797: 5785:. Retrieved 5734: 5730: 5698:. Retrieved 5687: 5678: 5666:. Retrieved 5658:The Guardian 5657: 5647: 5635:. Retrieved 5622: 5612: 5586: 5581: 5540: 5534: 5528: 5492: 5486: 5475: 5450: 5446: 5434: 5422:. Retrieved 5410: 5396: 5368: 5364: 5358: 5349: 5345: 5339: 5327:. Retrieved 5318: 5308: 5267: 5263: 5206: 5202: 5192: 5147: 5143: 5116:. Retrieved 5102: 5049: 5045: 4985: 4981: 4913: 4909: 4859: 4855: 4845: 4828: 4824: 4817: 4792: 4788: 4782: 4739: 4735: 4679: 4675: 4615: 4611: 4545: 4541: 4531: 4486: 4482: 4438: 4434: 4380: 4376: 4314: 4310: 4300: 4255: 4251: 4187: 4183: 4142: 4138: 4132: 4124:the original 4119: 4115: 4077: 4073: 4067: 4030: 4027:PLOS Biology 4026: 4016: 3976:(1): 19–30. 3973: 3969: 3963: 3920: 3916: 3889:. Retrieved 3874: 3866: 3836: 3832: 3822: 3813: 3773: 3769: 3759: 3745: 3705: 3701: 3691: 3682: 3642: 3638: 3628: 3619: 3579: 3575: 3565: 3530: 3526: 3516: 3481: 3477: 3467: 3424: 3420: 3358: 3354: 3290: 3286: 3226: 3222: 3186:. Retrieved 3166: 3162: 3151: 3138: 3131:. Retrieved 3122: 3112: 3069: 3065: 3033: 3023: 2988: 2978: 2966:. Retrieved 2955: 2948:Zimmer, Carl 2942: 2907: 2903: 2835: 2831: 2775: 2771: 2715: 2711: 2649: 2645: 2614:. Retrieved 2603: 2596:Zimmer, Carl 2554: 2550: 2540: 2504: 2498: 2473:. Retrieved 2464: 2455: 2412: 2408: 2401:Reich, David 2346: 2342: 2290: 2250: 2246: 2236: 2195: 2191: 2168:. Retrieved 2157: 2147: 2102: 2098: 2088: 2033: 2024: 2020: 2016: 2012: 2003: 1999: 1995: 1991: 1988: 1976:upregulates 1963: 1959: 1951: 1942: 1926: 1922: 1907: 1903: 1899: 1891: 1887: 1883:Wallace Line 1879: 1863: 1853:fifth finger 1791: 1787: 1782: 1776: 1768: 1753:, a partial 1749: 1742: 1731: 1691:interorbital 1684: 1677: 1648:parsimonious 1645: 1632:Neanderthals 1621: 1618: 1611: 1602:gyrification 1555: 1532: 1504: 1484: 1480: 1462:introgressed 1455: 1446: 1440:levels, and 1414: 1405: 1375: 1360: 1216: 1172: 1150:Ardipithecus 1149: 1068:Earlier apes 1025:Neanderthals 1009:Homo sapiens 1006: 1005: 992: 976: 968: 960: 950: 949: 935: 927: 917: 916: 902: 894: 886: 876: 875: 861: 846: 834:Ardipithecus 831: 830: 816: 808: 794: 781: 768: 758:Sivapithecus 755: 745:Oreopithecus 742: 729: 715: 707: 698: 685: 672: 660: 646: 633: 618: 447: 440: 437: 429: 425:hemizygosity 417:X chromosome 410: 401: 397:genetic load 389: 370:Analysis of 369: 345: 340:Neues Museum 300: 296: 294:(1.8–2.4%). 279: 266: 252:1000 Genomes 249: 232: 223:Svante Pääbo 198:Neanderthals 189: 185: 177: 153:Western Asia 146: 115: 88: 76:North Africa 65: 53:Neanderthals 40: 39: 29: 7761:Grandmother 7716:Shore-based 7677:Aquatic ape 7568:Tam Pa Ling 7463:H. ergaster 7282:P. robustus 7020:11 February 6565:(7): e105. 5759:10230/25596 5585:Dan Jones: 4742:(1): 6308. 4145:: 126–129. 3815:migrations. 2616:14 December 2557:(1): 1–11. 2475:21 November 2105:(8): 1163. 2044:Neanderthal 1933:Zhoukoudian 1893:entry into 1875:East Asians 1873:but not to 1866:Melanesians 1810:Lagar Velho 1745:Gravettians 1587:gray matter 971:H. ergaster 864:Ar. ramidus 849:Ar. kadabba 811:O. praegens 649:Pleistocene 478:This box: 348:Mezmaiskaya 336:Le Moustier 257:David Reich 227:Nobel Prize 173:South Asian 165:Philippines 134:Polynesians 130:Melanesians 118:East Asians 8005:Categories 7800:Skin color 7785:Bipedalism 7746:Killer ape 7558:Cro-Magnon 7457:Denisovans 7333:H. habilis 7297:Humans and 7182:A. ramidus 7175:A. kadabba 5352:: 232–234. 4383:(7): 358. 4116:Hypothesis 2968:31 January 2080:References 1978:hemoglobin 1870:Denisovans 1824:Denisovans 1759:Manot Cave 1674:Morphology 1658:bottleneck 1636:Denisovans 1496:cerebellum 1434:optic disk 1040:Denisovans 979:Au. sediba 953:H. erectus 920:H. habilis 710:Ou. turkae 411:There are 169:Andamanese 107:Ethiopians 57:Denisovans 47:and early 7919:Theorists 7886:Timelines 7766:Patriarch 7742:Behavior 7667:Gathering 7595:Ancestors 7340:H. naledi 7275:P. boisei 7246:A. sediba 6931:1312.7749 5957:206551893 5329:6 January 4923:1208.2238 4230:221882937 3747:component 3183:0960-9822 3133:25 August 2934:210955842 2228:210982481 2170:6 January 2121:2079-7737 2048:Denisovan 1845:Denisovan 1808:found at 1755:calvarium 1559:occipital 1491:phenotype 938:Au. garhi 662:Homininae 421:autosomes 376:El Sidrón 362:) or the 244:gene flow 237:of three 138:admixture 122:Europeans 105:(such as 7965:Category 7820:Language 7790:Skeleton 7485:H. longi 7239:A. garhi 7076:Hominins 7071:Taxonomy 7014:Archived 6991:30651539 6908:32128408 6844:32095519 6781:Archived 6764:32095519 6705:35197631 6648:31969706 6591:16895447 6540:21896735 6481:18944814 6473:22965354 6438:28938123 6389:25743789 6381:26449472 6335:22840920 6283:25043035 6221:23341637 6159:34388371 6102:29551270 6042:Archived 6019:22042846 5949:24136958 5906:21944045 5846:21940856 5781:Archived 5777:21179161 5694:Archived 5662:Archived 5631:Archived 5602:Archived 5597: ; 5573:26693109 5565:12802315 5519:16826514 5467:14022816 5415:Archived 5323:Archived 5292:25629628 5243:17452632 5184:14504393 5112:Archived 5108:"Oase 2" 5094:17085588 5076:30052409 5022:10377462 4960:23055938 4886:22513287 4809:29739306 4774:28740249 4706:28235201 4650:21868630 4582:20498832 4542:PLOS ONE 4523:17090677 4457:24336922 4409:30022013 4349:29033327 4292:23112810 4252:PLOS ONE 4222:32973032 4094:17027252 4059:15024415 4008:13581775 3955:26886800 3885:Archived 3863:23410836 3833:Genetics 3810:23082212 3770:PLOS ONE 3742:23082212 3702:PLOS ONE 3679:23082212 3639:PLOS ONE 3616:23082212 3576:PLOS ONE 3557:25683122 3508:25683119 3459:28102248 3393:24476815 3317:23410836 3287:Genetics 3261:22936568 3127:Archived 3104:23003860 3096:24476670 3015:38405782 3006:10888882 2962:Archived 2926:32004458 2870:28982794 2802:24532731 2772:Genetics 2750:24352235 2684:20448178 2610:Archived 2579:14329491 2531:32193295 2469:Archived 2447:27032491 2403:(2016). 2375:29852022 2287:27889059 2220:32001636 2164:Archived 2139:36009790 2058:See also 1910:Negritos 1829:Genetics 1816:and the 1814:Portugal 1471:selected 636:Pliocene 283:efficacy 203:Genetics 157:Oceanian 95:Cushitic 61:hominins 7977:Commons 7929:Fossils 7795:Muscles 7706:Cooking 7662:Hunting 7139:Orrorin 6982:6335398 6961:Bibcode 6899:7032934 6878:Bibcode 6835:7015685 6814:Bibcode 6755:7015685 6732:Bibcode 6696:8907066 6675:Bibcode 6639:8386425 6618:Bibcode 6582:1523253 6531:3174671 6508:Bibcode 6429:5679310 6351:Science 6326:3426505 6274:4134395 6253:Bibcode 6212:3568306 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Interbreeding between archaic and modern humans

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